Plant Taxonomy

PLANT TAXONOMY
Second Edition
ABOUT THE
AUTHOR
O P Sharma, with his 35 research articles published in national and international journals, 29
books written for university students, and 40 years experience of teaching, is an able researcher,
established Indian author, and an experienced teacher. His areas of research include pollen morphol-
ogy, angiosperm’s anatomy and mycology, with special focus on Indian Cyperaceae (with particular
interest on Cyperus). Over a dozen of Dr Sharma’s books have been published through internation-
ally known publishers like Tata McGraw-Hill Education Pvt. Ltd., and Macmillan India Ltd. He
has also revised Economic Botany, an internationally renowned text by late Professor Albert F Hill
(Harvard University, USA), a publication of McGraw-Hill, New York. Encouraging reviews of his
books have been published in reputed scientific journals, and his books on Practical Botany have
received appreciations from some eminent botanists including J D Dodge (England), T Christensen
(Denmark), J M Herr (USA) and C R Metcalfe (England).
Immediately after passing his MSc Botany in first division from C.C.S. University, Meerut and
thereafter PhD from the same university, Dr Sharma started his teaching career in 1967 as a faculty
member of Botany Department, Meerut College, Meerut, and retired from his active services as
a Reader from the same department in 2007. Besides attending several national and international
workshops, symposia and conferences during his four decades of teaching career, Dr Sharma is still
enjoying his post-retirement inning as an active author.
PLANT TAXONOMY
Second Edition
O P Sharma
Retired Reader
Department of Botany,
Meerut College, Meerut
Tata McGraw-Hill Education Private Limited

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CONTENTS
Preface to the Second Edition xvii
Preface to the First Edition xxi
1. Introduction 1
1.1 Taxonomy and Systematics: Synonyms or Independent Branches 1
1.2 Systematics and the Systematist 2
1.3 Objectives, Goals and Aims of Plant Systematics 3
1.4 Hierarchical Stages of a Systematist 4
1.5 Hierarchical Categories of Taxonomy 4
1.6 Basic Components of Taxonomy 4
1.7 Future of Plant Taxonomy 5
Test Your Understanding 6
Suggested Reading 6
2. History of Plant Taxonomy 8
2.1 History and Development of Plant Classification 8
2.2 Taxonomy—Our Contemporary 13
2.3 Chronological Development of Taxonomy in India 13
2.4 Some Indian Journals Related to Taxonomy 13
2.5 A Note on Botanical Survey of India 16
Suggested Reading 18
3. Classification 19
3.1 What is Classification? 19
3.2 Ranks of Plant Classification 19
vi Contents
3.3 Types of Systems of Classification 19

3.4 Some Important Systems of Classification 20
3.5 Comparison of Systems of Classification 35
3.6 Current Systems of Classification 35
4. Taxonomic Structure 46
4.1 Concept of Taxa 46
4.2 Concept of Species 47
4.3 Concept of Genus 51
4.4 Concept of Family 52
4.5 Taxa above Family Level 52
5. Plant Collection and Specimen Preparation 55
5.1 Which Type of Specimens should be Collected? 55
5.2 Field Equipment and Supplies 55
5.3 Organisation of the Field Press 57
5.4 How to Press Plant Specimens? 57
5.5 Drying of Specimens 57
5.6 Mounting of Specimens 58
5.7 Deep-freezing Methods 58
5.8 Labelling of Specimens 59
5.9 Identification of Specimens 60
6. Examination of a Plant Specimen 61
6.1 Equipment Needed for Examination of Plant Specimen 61
6.2 Instructions to be Followed while Examining a Plant Specimen 61
6.3 Guide to the Examination of Plant Specimen 62
7. Plant Identification 69
7.1 What is Identification? 69
7.2 Characters often Considered before Plant Identification 69
Contents vii
7.3 Identification with Keys 70

7.4 Some Unconventional Identification Methods 71
7.5 Artificial Keys for the Identification of 51 Common Families of Angiosperms 72
8. Plant Nomenclature, Botanical Names and Phylocode 78
(A) Plant Nomenclature 78
8.1 Fundamentals of Plant Nomenclature 78
8.2 Common Names and Scientific Names 79
8.3 Binomial Nomenclature 79
8.4 International Code of Botanical Nomenclature (ICBN) 80
8.5 Some Important Rules of Nomenclature 86
(B) Botanical Names 89
8.6 Common Prefixes used in Specific Epithets 89
8.7 Common Suffixes used in Specific Epithets 89
8.8 Plant Parts used as Epithets 90
8.9 Specific Epithets Linked with Colour 90
8.10 Specific Epithets Linked with Geography 90
8.11 Specific Epithets Linked with Size 90
8.12 Specific Epithets Linked with Habit 90
8.13 Specific Epithets Linked with Habitats 91
(C) Phylocode: A New System of Nomenclature 91
8.14 What is Phylocode? 91
8.15 Principles of the Phylocode 92
8.16 Phylocode: An Overview 93
8.17 Registration Database of Phylocode 93
8.18 Main Events in the History of Phylocode 93
8.19 Future of Phylocode 94
9. Modern Trends in Plant Taxonomy 96
9.1 External Morphology in Relation to Taxonomy 96
9.2 Vegetative Anatomy in Relation to Taxonomy 98
9.3 Floral Anatomy in Relation to Taxonomy 102
9.4 Cytology in Relation to Taxonomy or Cytotaxonomy 103
viii Contents
9.5 Palynology in Relation to Taxonomy 106

9.6 Embryology in Relation to Taxonomy 109
9.7 Chemistry in Relation to Taxonomy 111
9.8 Ecology in Relation to Taxonomy 112
9.9 Palaeobotany in Relation to Taxonomy 112
9.10 Electron Microscopy in Relation to Taxonomy 113
10. Numerical Taxonomy 115
10.1 What is Numerical Taxonomy? 115
10.2 Principles of Numerical Taxonomy 115
10.3 Logical Steps of Numerical Taxonomy 116
10.4 Advantages of Numerical Taxonomy 118
10.5 Applications of Numerical Taxonomy 119
11. Chemotaxonomy 121
11.1 What is Chemotaxonomy? 121
11.2 Purpose of Chemotaxonomy 121
11.3 A Brief History 121
11.4 Chemical Characters and their use in Taxonomy 122
12. Molecular Taxonomy 128
12.1 What is Molecular Taxonomy? 128
12.2 Basics used in the Techniques and Applications of Molecular Taxonomy 128
12.3 Names of Some Techniques used in Molecular Taxonomy 129
12.4 Relation of Chemotaxonomy and Molecular Taxonomy 129
12.5 Where do we Stand Today in Knowing DNA-sequencing Techniques and Why? 129
12.6 Status of Molecular Taxonomy in India 129
12.7 What does a Typical Molecular Systematic Analysis Require? 130
12.8 Assumptions and Uses of Molecular Taxonomy 130
12.9 Criticism and Future of Molecular Systematics 131
12.10 A Note on Molecular Markers 132
12.11 Some Specific Examples Showing Role of Molecular Systematics in Plants 132
Contents ix

13. Serotaxonomy 134
13.1 What is Serotaxonomy? 134
13.2 Some other Related Terms 134
13.3 General Features of Serological Reactions 135
13.4 Brief History of Serotaxonomy 135
13.5 General Process of Serotaxonomy 135
13.6 Examples of Serotaxonomic Importance 136
14. Phylogeny: Origin and Evolution of Angiosperms 138
14.1 What is Phylogeny? 138
14.2 What are Angiosperms? 138
14.3 General Principles of Angiosperm Phylogeny 139
14.4 Angiosperm Phylogeny: A Generally Accepted Picture 140
14.5 Monophyletic or Polyphyletic Origin 140
14.6 Fossil Records or Time of Angiosperms Origin 141
14.7 Probable Ancestors of Angiosperms: Some Theories 141
14.8 Primitive Angiosperms 143
14.9 Place of the Origin of Angiosperms 144
14.10 Origin of Monocots: Some Views 145
14.11 Lines of Evolution in Angiosperms 145
15. Botanical Library 148
15.1 Botanical Library and Systematics 148
15.2 Common Terms Used in a Botanical Library 148
15.3 Use of Botanical Library 149
15.4 Botanical Library as a Training Centre 149
15.5 Classification used in a Botanical Library 149
15.6 Major Botanical Libraries of the World 150
15.7 Future Information Systems 150
x Contents
16. Herbarium 152

16.1 Herbarium and its Limits 152
16.2 History of Herbarium Development 153
16.3 A Modern Herbarium 153
16.4 Twenty Major Herbaria of the World 154
16.5 Major Indian Herbaria 154
16.6 Functions of Herbaria 155
16.7 Precautions for Using Herbarium 156
17. Botanical Gardens 158
17.1 What are Botanical Gardens? 158
17.2 45 Major Botanical Gardens of the World: A Chronological Directory 158
17.3 Major Botanical Gardens of India 161
17.4 Largest Botanical Garden of India 162
17.5 Largest Botanical Garden of the World 163
17.6 Role of Botanical Gardens 163
18. Floral Formula and Floral Diagram 165
18.1 What is a Floral Formula? 165
18.2 Symbols Employed in Floral Formulae 165
18.3 What is a Floral Diagram? 166
18.4 What Does a Floral Diagram Inform? 167
18.5 How to Draw a Floral Diagram? 168
19. Position of Some Selected Families in Classification Systems
Proposed by Bentham and Hooker, Engler and Prantl,
Hutchinson, Takhtajan, Cronquist, and Thorne 171
20. 550 Terms of Plant Description 176
20.1 Plant Parts 176
20.2 Plant Types (Habit and Habitat) 176
Contents xi
20.3 Root 177

20.4 Stem (Types, Surface, Forms and Modifications) 179
20.5 Leaf 182
20.6 Inflorescence 192
20.7 Flower (Parts and Types) 194
20.8 Calyx 196
20.9 Aestivation 196
20.10 Corolla 197
20.11 Perianth 199
20.12 Androecium 199
20.13 Gynoecium 202
20.14 Placentation 205
20.15 Fruit 206
21. Selected Families of Dicotyledons 212
21.1 Dicotyledons 212
21.2 Polypetalae 213
21.3 Ranales 213
21.4 Magnoliaceae (Magnolia Family) 214
21.5 Annonaceae (Custard Apple Family) 217
21.6 Ranunculaceae (Buttercup Family) 221
21.7 Nymphaeaceae (Water-Lily Family) 227
21.8 Parietales 231
21.9 Papaveraceae (Poppy Family) 231
21.10 Fumariaceae (Fumaria Family or Fumitory Family) 236
21.11 Capparidaceae (The Capers Family) 239
21.12 Brassicaceae or Cruciferae (Mustard Family) 243
21.13 Violaceae (Violet Family) 246
21.14 Caryophyllineae 249
21.15 Caryophyllaceae (Pink Family) 249
21.16 Portulacaceae (Purslane Family) 253
21.17 Malvales 257
21.18 Malvaceae (Mallow Family) 257
21.19 Sterculiaceae (Sterculia Family) 262
21.20 Tiliaceae (Lime Family or Basswood Family) 265
xii Contents
21.21 Bombacaceae (Bombax Family) 268

21.22 Geraniales 271
21.23 Geraniaceae (Geranium Family) 271
21.24 Oxalidaceae (Wood-Sorrel Family) 274
21.25 Rutaceae (Citrus Family or Rue Family) 276
21.26 Meliaceae (Mahogany Family) 281
21.27 Celastrales 284
21.28 Rhamnaceae (Buckthorn Family) 285
21.29 Vitaceae (Grape Family) 286
21.30 Sapindales 289
21.31 Sapindaceae (Soapberry Family) 289
21.32 Anacardiaceae (Cashew Family) 293
21.33 Rosales 297
21.34 Leguminosae (Legume Family) 297
21.35 Subfamily Mimosoideae or Mimoseae (Mimosa Family) 299
21.36 Subfamily Caesalpinioideae or Caesalpinieae (Cassia Family) 302
21.37 Subfamily Papilionoideae (Pea Family or Bean Family) 306
21.38 Rosaceae (Rose Family) 312
21.39 Saxifragaceae (Saxifrage Family) 318
21.40 Myrtales 320
21.41 Combretaceae (Combretum Family) 321
21.42 Myrtaceae (Myrtle Family) 323
21.43 Lythraceae (Loosestrife Family) 327
21.44 Passiflorales 329
21.45 Cucurbitaceae (Gourd Family) 329
21.46 Begoniaceae (Begonia Family) 333
21.47 Passifloraceae (Passion-Flower Family) 336
21.48 Ficoidales 338
21.49 Cactaceae (Cactus Family) 339
21.50 Umbellales 341
21.51 Umbelliferae or Apiaceae (Carrot Family) 342
21.52 Araliaceae (Aralia or Ginseng Family) 345
21.53 Gamopetalae 348
21.54 Rubiales 349
21.55 Rubiaceae (Coffee Family or Madder Family) 349
21.56 Caprifoliaceae (Honeysuckle Family) 352
Contents xiii
21.57 Asterales 355

21.58 Compositae or Asteraceae (Sunflower or Aster Family) 355
21.59 Campanales 362
21.60 Campanulaceae (Bellflower Family) 362
21.61 Ericales 365
21.62 Ericaceae (Heath Family) 365
21.63 Primulales 368
21.64 Primulaceae (Primula or Primrose Family) 369
21.65 Plumbaginaceae (Leadwort Family) 372
21.66 Ebenales 374
21.67 Sapotaceae (Sapota Family) 375
21.68 Gentianales 377
21.69 Oleaceae (Olive Family) 377
21.70 Asclepiadaceae (Milkweed Family) 380
21.71 Apocynaceae (Dogbane Family) 384
21.72 Loganiaceae (Logania Family) 389
21.73 Polemoniales 391
21.74 Boraginaceae (Borage Family) 391
21.75 Convolvulaceae (Morning Glory Family) 394
21.76 Solanaceae (Potato or Nightshade Family) 397
21.77 Polemoniaceae (Phlox Family) 401
21.78 Personales 405
21.79 Scrophulariaceae (Snapdragon Family) 405
21.80 Bignoniaceae (Bignonia Family) 409
21.81 Pedaliaceae (Benne Family) 411
21.82 Acanthaceae (Acanthus Family) 414
21.83 Lamiales 417
21.84 Labiatae or Lamiaceae (Mint Family) 419
21.85 Verbenaceae (Verbena Family) 423
21.86 Plantaginaceae (Plantago Family) 425
21.87 Monochlamydeae 428
21.88 Chenopodiaceae (Goosefoot Family) 429
21.89 Amaranthaceae (Amaranthus or Pigweed Family) 432
21.90 Polygonaceae (Buckwheat or Smartweed Family) 435
21.91 Aristolochiaceae (Birthwort Family) 438
21.92 Piperaceae (Pepper Family) 440
xiv Contents
21.93 Loranthaceae (Mistletoe Family) 443

21.94 Euphorbiaceae (Spurge Family) 445
21.95 Urticaceae (Nettle Family) 451
21.96 Moraceae (Mulberry Family) 453
21.97 Cannabinaceae (Hemp Family) 457
21.98 Casuarinaceae (Casuarina Family) 460
21.99 Salicaceae (Willow Family) 463
22. Selected Families of Monocotyledons 469
22.1 Monocotyledons and their Classification 469
22.2 Orchidaceae (Orchid Family) 470
22.3 Iridaceae (Iris Family) 474
22.4 Amaryllidaceae (Daffodil Family) 477
22.5 Bromeliaceae (Pineapple Family) 479
22.6 Cannaceae (Canna Family) 482
22.7 Musaceae (Banana Family) 485
22.8 Zingiberaceae (Ginger Family) 488
22.9 Liliaceae (Lily Family) 490
22.10 Commelinaceae (Spiderwort Family) 494
22.11 Juncaceae (Rush Family) 497
22.12 Palmae or Arecaceae (Palm Family) 500
22.13 Typhaceae (Cattail Family) 504
22.14 Araceae (Arum Family) 507
22.15 Alismataceae or Alismaceae (Water Plaintain Family) 509
22.16 Cyperaceae (Sedge Family) 513
22.17 Gramineae or Poaceae (Grass Family) 518
Appendix 1 526
Examination Tools—Comparative Tables of Selected Families 526
Table A.1 Magnoliaceae and Annonaceae 526
Table A.2 Ranunculaceae and Nymphaeaceae 526
Table A.3 Ranunculaceae and Cruciferae (Brassicaceae) 527
Table A.4 Ranunculaceae and Rosaceae 527
Table A.5 Papaveraceae and Fumariaceae 528
Contents xv
Table A.6 Papaveraceae and Cruciferae (Brassicaceae) 528

Table A.7 Capparidaceae and Cruciferae (Brassicaceae) 528
Table A.8 Capparidaceae and Caryophyllaceae 529
Table A.9 Malvaceae and Tiliaceae 529
Table A.10 Oxalidaceae and Geraniaceae 529
Table A.11 Rutaceae and Meliaceae 530
Table A.12 Rutaceae and Anacardiaceae 530
Table A.13 Sapindaceae and Anacardiaceae 531
Table A.14 Mimosoideae, Caesalpinioideae and Papilionoideae 531
Table A.15 Rosaceae and Myrtaceae 531
Table A.16 Rosaceae and Cucurbitaceae 532
Table A.17 Rosaceae and Saxifragaceae 532
Table A.18 Umbelliferae and Rubiaceae 533
Table A.19 Umbelliferae (Apiaceae) and Compositae (Asteraceae) 533
Table A.20 Rubiaceae and Compositae (Asteraceae) 534
Table A.21 Asclepiadaceae and Apocynaceae 534
Table A.22 Convolvulaceae and Solanaceae 534
Table A.23 Labiatae, Boraginaceae and Scrophulariaceae 535
Table A.24 Acanthaceae, Labiatae and Verbenaceae 535
Table A.25 Chenopodiaceae and Amaranthaceae 536
Table A.26 Amaranthaceae and Polygonaceae 536
Table A.27 Euphorbiaceae and Moraceae 537
Table A.28 Musaceae and Zingiberaceae 537
Table A.29 Liliaceae and Iridaceae 538
Table A.30 Liliaceae and Amaryllidaceae 538
Table A.31 Juncaceae and Cyperaceae 538
Table A.32 Gramineae (Poaceae) and Juncaceae 539
Table A.33 Cyperaceae and Gramineae (Poaceae) 539
Examination Tools—Major Characters of Discussed Families 540
Appendix 2 547
Examination Tools—Selected Medicinal Plants and their Utility: At a Glance 547
Index 553
PREFACE TO THE
SECOND EDITION
Plant Taxonomy has been revised to reflect global perspectives with particular emphasis on the
current syllabi requirements of the Indian Universities. Published first in 1993 and reprinted 19 times
in subsequent years upto 2008, speaks volumes about its overwhelming reception by the readers. The
developments in the field of plant systematics, coupled with new and exciting discoveries contributing
to the progress in taxonomical fields providing new information in classification of plants, have
given impetus to bring out the revised edition. The new edition was also needed because of several
changes made in the syllabi of Indian Universities, and also because of variations in the market
trends and examination patterns.
The second edition of Plant Taxonomy is designed to present the current principles, practices
and techniques of plant taxonomy and contemporary classifications, and also to describe some of the
other important angiospermic families and groups. Those who are familiar with the first edition will
be aware that there has been change of format to a larger page size, and also a completely different
layout of the text. A considerable number of illustrations and chapter-wise revision questions have
also been added in the revised text.
The major aim of this book is to provide a general overview of plant taxonomy in the most effective
and positive manner to the students. This revised edition of Plant Taxonomy is a broad, up-to-date
synthesis of this active and fascinating field of botany. Besides retaining all chapters of the first edition,
this revised edition now also includes some new chapters, viz. Taxonomic Structure, Examination of
Plant Specimen, Molecular Taxonomy, and Table of Major Characters of Discussed Families. Also
added in this new edition are the detailed discussions of 16 more families of dicotyledons (viz.
Saxifragaceae, Begoniaceae, Passifloraceae, Araliaceae, Caprifoliaceae, Campanulaceae, Ericaceae,
Plumbaginaceae, Loganiaceae, Polemoniaceae, Pedaliaceae, Plantaginaceae, Aristolochiaceae,
Piperaceae, Cannabinaceae, and Salicaceae) and 4 families of monocotyledons (viz. Bromeliaceae,
Cannaceae, Juncaceae, and Typhaceae), thus increasing the number of discussed families from 69
to 89 out of total number of 200 families of flowering plants discussed by Bentham and Hooker.
This number of discussed families in this book is now larger than all other available books on plant
taxonomy in the Indian market. The revised edition also discusses some new subtopics in certain
existing chapters as suggested by reviewers of the book. These subtopics include (i) Deep-freezing
methods, (ii) Restructuring of ICBN with the addition of the details of St. Louis Code (1999) and
xviii Preface to the Second Edition
Vienna Code (2005), (iii) Phylocode, a new system of nomenclature, and (iv) Classification and major
characters of subclasses and orders of dicotyledons and monocotyledons. In Comparative Tables
of Selected Families, 13 new tables have been added, thus increasing the total number from 20 to
33 comparative tables. Another new feature is an index-listing of all relevant medicinal usages of
discussed plants given at the end of the book. These additions will prove to be of great help to the
students while preparing for their examination.
The major highlights of this book are as follows:
• Complete coverage of all important topics in Plant Taxonomy
– Plant Classification
– Plant Collection and Specimen Preparation
– Identification and Botanical Nomenclature
– Herbarium and Botanical Gardens
– Phylogeny
– Classification System and Description of 89 Families
• Four recent application-based Chapters
– Numerical Taxonomy
– Chemotaxonomy
– Serotaxonomy
– Molecular Taxonomy
• Examination Preparation Tools to aid students memorise basics and prepare efficiently for
their examinations
– All 89 Families summarised in a Table
– 33 questions on ‘Differentiate between the families’
– 164 Chapter-end Examination Review Questions
• Rich pedagogy
– More than 900 rich plant organs sketches and floral diagrams
– Around 40 tables
• Updated as per International Botanical Congress Guidelines, (St. Louis, Missouri, July–
August 1999) organised by IAPT: St. Louis Code, The International Code of Botanical
Nomenclature.
Plant Taxonomy, in its present form, should now meet the needs of undergraduate students of
all universities of the country taking botany as their major subject. It should also cater to the com-
plete requirements of postgraduate students of Botany, Agriculture and Forestry in majority of the
Indian Universities. As has already been in the past, the book should also be useful for and liked
by students preparing for AIPMT, CPMT, NET, SLET, IAS, IFS, PCS and several other major
competitive examinations.
A major highlight of this revised edition is that all chapters are largely independent, so that the
teacher may choose the desired sequence of topics according to his/her syllabus requirements. The
level of presentation is primarily for undergraduate students and based mainly on the assumption
that the students will have had an introductory Botany course. The author hopes that the book in its
Preface to the Second Edition xix
present form will prove stimulating to serious amateurs, teachers as well as professionals who are
specialists in other fields but use classifications and other taxonomic information about flowering
plants.
Before going to press, the revised manuscript was critically reviewed by some experts in the field,
including Dr Tariq Husain, Angiosperm Taxonomy and Herbarium Division, National Botanical
Research Institute, Lucknow; Dr N K Soni, Department of Botany, Dr H S Gour University, Sagar;
and Dr P J Handique, Department of Biotechnology, Gauhati University, Guwahati. Besides several
positive comments and suggestions of these reviewers, one of them even commented that, “This book
is above all the books available on Plant Taxonomy in the country”. For the pains taken by these
reviewers, the author acknowledges their help and thank them all by heart.
Without the help of my dear student, Dr Mayank Uday Charaya, Professor of Botany, C.C.S.
University, Meerut, this book would have not come in its present form. I express my heartfelt feelings
to Professor Charaya and his team for searching latest information through the Internet. For several
types of suggestions, technical clarifications and encouragements, I also express my gratitude to Dr
R Shiam (my teacher from BSc to PhD), Dr N P Saxena (Meerut), Dr Lokendra Singh (Meerut)
and Dr H P Pandey (University of Allahabad).
At this stage, I cannot ignore the help, support, cooperation and total dedication that my wife
Dr (Mrs) Kanti D Sharma, PhD extended to me during the entire period of the preparation of this
book. She deserves all my love and appreciation. Throughout this period, I had no option but to
ignore my grandchildren (Kuhu and Karan), though deeply unwillingly. To them, I can simply say
that I love both of you to my fullest.
I would be grateful if users of the book notify me of any errors or omissions that come to their
notice. Comments and suggestions for improvements may be sent to tmh.sciencemathsfeedback@
gmail.com (kindly mention the title and author name in the subject line).
Jun 2009 O P Sharma

+91-9837566555
PREFACE TO THE
FIRST EDITION
As opposed to the various other branches of Botany, Plant Taxonomy, has for a long time lacked
an appropriate text, especially at the undergraduate level. Currently a student can choose between
the voluminous reference books meant for advanced and the much abbreviated college texts. I have
tried an approach that would interest and inspire all those who take up Plant Taxonomy. The aim
of this book is to provide a general overview of Plant Taxonomy in an effective manner—a simpli-
fied but factual account.
My teaching experience of over 25 years has convinced me that a study of the basic principles of
Plant Taxonomy and of selected families of angiosperms provide sufficient material for a beginning
course on this subject at the undergraduate level. Consequently, I have discussed the basic principles of
the discipline and then moved on to some selected families of dicotyledons and monocotyledons.
Botanists agree that presently there are about 2,45,000 species of flowering plants on earth and
these have been grouped into 197–430 families, depending on the authority concerned. However,
a basic knowledge of angiosperms can be acquired by a study of a selected group of families. I
have selected 69 families of flowering plants, prominent in India. The criteria for selection are easy
availability of their several ‘representatives’ unusual characteristics, economic importance, typical
representation of a particular taxa.
In limiting my discussion to 69 families, I have had to deliberately overlook many which are
important and commonly available in several parts of the world except the Indian subcontinent.
While arranging the families I have followed Bentham and Hooker’s system since (i) It is followed
in almost all Indian herbaria, (ii) Majority of the Indian flora are based on this system, (iii) It is
the most appropriate system for studying angiosperms in laboratories.
The subject matter is divided into twenty chapters which are by-and-large independent so that
the teacher may choose the desired sequence of topics. Chapters 1–3 discuss the traditional aspects
of Plant Taxonomy including the history and classification. Chapters 4–6 discuss the methods,
techniques and principles of plant collection, Plant identification and nomenclature. Chapter 7
discusses the role of various Botanical disciplines—External Morphology, Vegetative Anatomy,
Floral Anatomy, Cytology, Palynology, Embryology, Chemistry, Ecology, Palaeobotany, Electron
Microscopy—in solving various taxonomic problems.
xxii Preface to the First Edition
Chapters 8–10 deal with Numerical Taxonomy, Chemotaxonomy and Serotaxonomy, respectively.
Latest information on phylogeny of angiosperms, the various theories regarding their origin and evolu-
tion, is presented in Chapter 11. Chapters 12–14 expand on various botanical institutions—botanical
libraries, herbaria and botanical gardens.
Floral formulae and floral diagrams, from which it is generally possible to identify a family, are
explained in Chapter 15. In Chapter 16, position is assigned to 72 commonly available families in
some well-known systems of classification. Chapter 17 discusses over 550 common botanical terms
used in taxonomic description.
Chapters 18 and 19 provide descriptions, illustrations and related information on 69 families
of flowering plants—57 dicotyledons and 12 monocotyledons. Description of each family includes its
systematic position, filed recognition, distribution, general characteristics, pollination and dispersal,
general floral formula, economic importance, systematics and phylogeny, and also the descriptions
of some common plants chosen as typical representatives of the family. Chapter 20 contains twenty
tables of comparison between different families.
Although this book is primarily for undergraduate students of universities in the Indian subcon-
tinent, I am also hopeful that it will benefit postgraduates, teachers and professionals.
I would be grateful if the reader notifies me of any errors or omission. I also welcome comments
and suggestions from students and fellow teachers.
May, 1993 O P Sharma

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INTRODUCTION 1
1.1 TAXONOMY AND SYSTEMATICS: SYNONYMS OR
INDEPENDENT BRANCHES
The “science of the classification of organisms according to their resemblances and differences” is
the definition of taxonomy as stated by Abercrombie et al. (1980) in the 7th edition of The Penguin
Dictionary of Biology. According to them the word systematics is “often used synonymously with
taxonomy, but sometimes interpreted more widely to include also the identification, practice of clas-
sification and nomenclature”. However, Andrew Sugden in the 1986 edition of Longman’s Illustrated
Dictionary of Botany defined taxonomy as “the science of classification and relationships of organ-
isms”, and systematics as “the part of classification that involves the arrangement of organisms into
related groups”.
Jones and Luchsinger (1987) believe that although there is no agreement among the botanists
“for the distinctions between systematics and taxonomy”, some botanists treat them as two separate
branches. According to these botanists, systematics is “the study of the diversity of plants and their
identification, naming, classification and evolution” while taxonomy is “restricted to the study of
classification” (Jones and Luchsinger, 1987).
Solbrig (1966) maintained that the two fields of inquiry (viz. taxonomy and systematics) have
absolutely different concern. But Heywood (1967) and Ross (1974) treated systematics as a field cov-
ering the scientific study of the diversity and differentiation of organisms and the relationships that
exist between them, and consider taxonomy as a part of systematics. Morse (1974), while presenting
a system of computer programming to help in plant identification, treated taxonomy and systematics
as two separate branches. Stace (1980) also considered these two as separate disciplines in his book
entitled Plant Taxonomy and Biosystematics.
However, the terms taxonomy and systematics have been so loosely and interchangeably used
in the past that to establish a proper delineation between the two is extremely difficult. In actual
practice, and also in the present text, the two terms are used synonymously and deal with the study
of classification, its principles, procedures and rules. Lam (1959) and Turrill (1964) also used them
as synonyms.
2 Plant Taxonomy
The Greek meaning of taxonomy is “arrangement by rules” and of systematics is “to put together”.
The term taxonomy was first coined and used by the famous French botanist A.P. de Candolle in
1813 in his book Theorie Elementaire de la Botanique.
1.2 SYSTEMATICS AND THE SYSTEMATIST

1.2.1 Systematics
Radford (1986) defined systematics as “the study of phenotypic, genetic and phylogenetic relationship
among taxa”, and mentioned that the science of systematics possesses following essential qualities:
From the fundamental point of view systematics is the study of the nature, causes, patterns, and
trends in variation among taxa.
From the structural point of view systematics is the study of the basic taxonomic components and
evolution.
From the functional point of view systematics is the study of characters from many fields of
evidence for establishing relationships among taxa.
From the developmental point of view systematics is the study of variation among taxa for the
determination of character correlations and relationships.
From the theoretical point of view systematics is based on the idea that because of great diver-
sity in the biological world there exist some ‘discontinuous units’ that can be identified, classified,
described and named on the basis of evolution.
From the philosophical point of view systematics is an ever-evolving and unending field for
understanding taxonomic and evolutionary processes, principles and concepts.
1.2.2 Systematist
The systematist is a student, researcher, and scholar who studies, classifies, identifies, describes,
names, observes, synthesizes or analyses the variations within the populations, species and higher
taxa.
From the functional standpoint, a taxonomist has the following activities or duties to perform:
1. As a classifier, a taxonomist determines the position and rank for new taxa.
2. As an identifier, a taxonomist distinguishes or identifies new taxa and establishes the diag-
nostic characters for old taxa.
3. As a describer, a taxonomist determines the circumscription for a specimen according to an
established system of classification.
4. As a nomenclaturist, a taxonomist assigns names to new taxa and determines the correct
names for old taxa according to the rules laid down by the latest International Code of
Botanical Nomenclature.
5. As a coiner, a taxonomist selects the proper Greek or Latin prefixes or suffixes for scientific
names.
Introduction 3
6. As an observer, a taxonomist observes the characteristics of organisms, population and taxa

for a better understanding of their taxonomy.
7. As an analyzer, a taxonomist decides which characters of the individual, population or taxa
are to be analyzed.
8. As a synthesizer, a taxonomist collects information from various aspects (morphology, cytol-
ogy, anatomy, etc.) of the organisms for the development of new systems of classification and
identification.
9. As a theorizer, a taxonomist develops new concepts on the basis of recent findings and
reinterprets old concepts.
10. As a historian, a taxonomist provides better understanding of the old classical concepts in
the light of the new researches in the field.
11. As a generator, a taxonomist generates effective methods for collection, analysis and presen-
tation of data.
12. As a conservator, a taxonomist suggests new approaches for the preservation of threatened
and endangered species.
1.2.3 Every Human Being is a Taxonomist

Every human being always behaves as a taxonomist. It is so because each one of us identifies, clas-
sifies, describes, and names while taking decisions about the food we eat, the beverages we drink,
the homes we purchase, the clothes we wear, the friends we like, the games we play, the religion we
believe in, and the politics we exercise. While talking about sex, sickness, sin, and sorrow we talk
differently to children, old people, specialists, and parents, and thus behave as a taxonomist in the
daily routine life. By smelling we identify garlic, by tasting we recognise sugar, and by seeing we
name the animal as elephant, and thus behave as taxonomists. We describe that road is long, grass
is green and ball is round, and thus work as a taxonomist.
1.3 OBJECTIVES, GOALS AND AIMS OF PLANT SYSTEMATICS

1.3.1 Objectives
1. To prepare a scheme of classification that provides phenetic, natural or phylogenetic relation-
ships among plants.
2. To establish a suitable method for identification, nomenclature and description of plant
taxa.
3. To provide an inventory of plant taxa that suits local, regional, and continental needs.
4. To create an understanding of the evolutionary processes.
5. To train the students of plant sciences in regard to the diversity of organisms and their rela-
tionship with other biological branches.
1.3.2 Goals
1. To acquire the fundamental values of plants systematics.
2. To know about the basic concepts and principles of plant systematics.
4 Plant Taxonomy
3. To be aware of the importance of taxonomic relationships in plant systematics.

4. To develop the knowledge of applicability of plant systematic studies.
1.3.3 Aims for the Study of Regional Flora

1. To know how to collect specimens.
2. To know how to prepare specimens for future preservation.
3. To know how a manual should be used.
4. To know how to use identification keys.
5. To recognize divisions, classes, orders, families, genera, and species.
6. To know how the plants are described.
7. To know how the diversity in species may be related with the regional habitat diversity.
8. To become familiar with the basic taxonomic principles, and with at least one system of
plant classification.
1.4 HIERARCHICAL STAGES OF A SYSTEMATIST

The following may be the hierarchical grades or stages of a systematist:
A natural scientist is a systematist who provides or gathers information about the relationships
between phenetic, genetic, and ecological variations among the organisms.
A systematist is a taxonomist who develops a profound knowledge of the specific aspects of
systematics.
A taxonomist is a classifier who names, describes, classifies and identifies the organisms.
A classifier is a student who establishes the named, delimited, and identified taxa for establishing
the diversity among organisms.
A student of diversity among organisms is a classifier, taxonomist, systematist and natural sci-
entist who provides information to the society about the phenetic, genetic, and ecological variations
among the organisms for the upliftment of mankind.
1.5 HIERARCHICAL CATEGORIES OF TAXONOMY

The International Code of Botanical Nomenclature (Voss et al. 1983) has recognized the following
categories or ranks of taxa along with their endings, as mentioned in Table 1.1.
1.6 BASIC COMPONENTS OF TAXONOMY

Classification, identification, description and nomenclature are the four basic components of
taxonomy.
Classification is the arrangement of botanical groups with definite circumscriptions by position
and rank according to artificial criteria, phenetic similarities, or phylogenetic relationships.
Identification is the determination of similarities or dissimilarities between the two elements.
Under identification we make a direct comparison of the characteristic features of a specimen with
those present in the already existing keys for identification.
Introduction 5
Table 1.1 Series of ranks and endings provided by the International Code of Botanical Nomenclature
(1983)
Ranks of Taxa Endings of ranks above genus Examples
Division phyta Pterophyta, Magnoliophyta

Subdivision phytina Pterophytina
Class opsida Pteropsida, Magnoliopsida
Subclass opsidae Pteropsidae
Order ales Rosales, Asterales
Suborder ineae Rosineae
Family aceae Rosaceae, Asteraceae
Subfamily oideae Rosoideae
Tribe eae Roseae
Subtribe inae Rosinae
Genus us, a, um, on, es, etc. Rosa, Pinus
Subgenus
Section
Subsection
Series
Subseries
Species
Subspecies
Variety
Subvariety
Form
Subform
Description is the orderly recording of maximum possible characters of a taxon, individual plant,
plant part, or object.
Nomenclature is a simple system under which the individual taxonomic groups of plants are
scientifically named.
1.7 FUTURE OF PLANT TAXONOMY

The evidence gathered from written records indicates that man has been categorizing or classify-
ing organisms for more than 2000 years. Incorporation of new researches, evidences, methods and
techniques have developed the science of systematics to its present status. But a lot more is still to
be done. Introduction of mathematical data, statistical details and the use of computers in the recent
past have brought tremendous changes in the field of taxonomy. The use of electron microscopy in
several cases has entirely changed the old concepts. Chromatography and spectrochemistry have also
brought unexpected changes in several existing ideas.
In the future, several new aspects in the field of systematics, will be explored. Some of them are
undermentioned:
6 Plant Taxonomy
1. According to Stace (1980) about 3,00,000 species of green plants, over 1,50,000 fungi and
a few thousand bacteria are known to the biologists. But in future several thousands of
new species of plants still await discovery, description, and naming in different parts of the
world.
2. Monographs of many plant groups are to be prepared.
3. Many groups, treated more than a century ago, badly need a revision in the light of new
discoveries and techniques.
4. Explorations of several unknown floristic regions are highly essential.
5. Several old floras are to be revised.
6. Speciation understanding in several groups is yet to be done.
7. Many old basic fields and taxonomic principles need re-evaluation.
8. Biology of a majority of the plant species has not been studied so far.
9. Several mathematical and statistical schemes for the determination of relationships have been
developed. These schemes are yet to be used in many cases. More mathematical logic is to
be used in taxonomic investigations.
10. Several ‘anomalous groups’ are still waiting for their proper placing in the systems of
classification.
11. In future, taxonomists will have to use the expertise and knowledge of genetic diversity. This
would help in the introduction of new crops and also in the improvement of old crops.
12. New and improved methods of studying nucleic acids will have to be used by the taxonomists
in future for the basic understanding of the relationships among the groups of organisms.
Test Your Understanding

1. Taxonomy and Systematics: Are they synonyms or two independent branches?
2. Enlist any six functions, a taxonomist has to perform.
3. Write a brief note on the fact that every human being is a taxonomist.
4. International Code of Botanical Nomenclature has recognised some basic categories or ranks
of taxa. Name any five of them along with their endings and an example of each of them.
5. Describe the future of plant taxonomy in about 100 words.
Suggested Reading
Bell, C.R., 1969, Plant Variation and Classification, Macmillan, London.
Davis, P.H. and V.H. Heywood, 1963, Principles of Angiosperm Taxonomy, Oliver & Boyd, Edinburgh.
Heywood, V.H., 1967, Plant Taxonomy, Edward Arnold Publishers, London.
Jones, S.B. and A.E. Luchsinger, 1987, Plant Systematics (2nd Ed.), McGraw-Hill, New York.
Lam, H.J., 1959, ‘Taxonomy: General principles and angiosperms,’ Vistas in Botany 2: 4–75.
Introduction 7
Morse, L.E., 1974, ‘Computer assisted storage and retrieval of the data of taxonomy and systematics,’
Taxon 23: 29–43.
Radford, A.E., 1986, Fundamentals of Plant Systematics, Harper and Row, New York.
Stare, C.A., 1980, Plant Taxonomy and Biosystematics, University Park Press, Baltimore.
Voss, E.G., 1983, International Code of Botanical Nomenclature, Bohn, Scheltema and Holkema, Utrecht,
Netherlands.
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HISTORY OF
PLANT TAXONOMY 2
2.1 HISTORY AND DEVELOPMENT OF PLANT CLASSIFICATION
2.1.1 Earlier than Man Could Read and Write
Not much is known today about the botanical knowledge of our preliterate ancestors. But it is clear
that they knew by experience the plants that were edible and others which were not. The preliterate
mankind also possessed some kinds of linguistic mechanisms for correct distinctions among differ-
ent kinds of plants.
2.1.2 Age of Theophrastus, Secundus, Dioscorides and Parasara

The advent of writing and printing changed the entire picture of the ancient preliterate mankind.
Theophrastus (370–285 B.C.), the “grandfather of the modern botany”, was the greatest botanical
writer of the distant past. He was a student of Plato and Aristotle, and studied botany under their
philosophic guidance at Athens. He classified the plants into four groups: herbs, subshrubs, shrubs,
and trees. He also distinguished between the nonflowering plants (Cryptogams) and flowering plants
(Phanerogams). He suggested that calyx and corolla are the modified leaves. He described nearly 500
plants in details, and certain names (e.g. Asparagus, Daucas, and Narcissus) are used even today
in the same sense. The details of his works are available to the world in the form of books entitled
“Enquiry into Plants” and “The Causes of Plants”.
Caius Plinius Secundus (23–79 A.D.), known to the botanical world as Pliny the Elder, was a
lawyer and also served in the Roman army till his death caused by the heart failure. Voluminous and
important works of Pliny are available in the form of 37 volumes of Natural History. He described
the biological, medicinal and agricultural aspects of the plants known to the world up to his time in
these volumes, and Pliny’s Natural History was among the first books to be printed by the movable
type in the late 15th century. The word ‘stamen’ in its modern sense was first used by Pliny.
Pedanios Dioscorides (62–128 A.D.) was a contemporary of Pliny the Elder. He was actually a
Physician in the Roman army and belonged to a Roman province, Cilicia. His monumental work is
compiled in the form of Materia Medica. It described the botany, mainly the medicinal aspects of
about 600 species of plants. Materia Medica was considered as a highly esteemed source book for
History of Plant Taxonomy 9
those practicing medicine in those days. The importance and fame of Materia Medica of Dioscorides
may be guessed from the fact that the Emperor Flavius Anicius Olybrius (500 A.D.) presented a
beautifully illustrated copy of the book to his daughter (Princess Juliana Anicia) as a precious
gift at the time of her marriage. Juliana’s copy of Materia Medica remained for several years in
Constantinople (Istanbul) and was later transferred to Vienna, where it still exists under the name
of Codex Juliana. Several plant names (e.g. Aloe, Aristolochia, Anemone, Phaseolus) as suggested
by Dioscorides are still in use in the present botanical literature.
The book titled Vrikshayurveda, written by an Indian, Parasara, is one of the earliest Indian
works describing plants in a scientific manner. Though it was written before the beginning of the
Christian era, the scientifically described plant classification and distribution in this book led the
famous systematist Albert E. Radford (1986) of USA to state that Parasara had some kind of hand
lens or microscope. There exist several other references which show that the early civilizations of
India, Egypt and China did have a definite knowledge of the plant taxonomy.
2.1.3 Taxonomy in Middle Ages

Little is known about the development of taxonomy during the early one thousand years of the
Christian era. Then came the Middle Ages or Medieval Ages (about A.D. 1100–1500). Albertus
Magnus (A.D. 1200–1280) has been the most famous plant taxonomy worker of this period. Commonly
called “Doctor Universalis” or “Aristotle of the Middle Ages” by his contemporaries and historians,
Magnus employed a scheme of classification of plants that recognised monocots and dicots, and
separated nonvascular plants from vascular plants. Worthmentioning are the names of two Muslim
scholars of twelfth century. Ibn-Sina who authored Canon of Medicine, and Ibn-al-Awwam of Spain
who described over 600 species of plants and interpreted accurately the sexuality in plants.
2.1.4 Herbalists
After the Medieval Ages (A.D. 1500) the history of plant taxonomy was influenced tremendously by
two things, the invention of printing and the development of the science of navigation. The printing
technology lowered the cost of books and increased literacy. During the early years of printing, the
medically oriented books on plants became quite popular. Printed forms of ancient texts had many
superfluous and irrelevant writings and this actually prompted several interested persons to write
and publish their own botanical medical books. These books were called herbals and their authors
were called herbalists.
The science of navigation prompted sailors to go on long voyages. This resulted in the explora-
tion of several new areas of the world, and, in turn, increased man’s practical knowledge of plant
taxonomy.
At the advent of 16th century the first herbals published were “Gart der Gesundheit” and “Hortus
Sanitalis”. These herbals had crude illustrations of plants and were published without an attribution of
authorship. However, the 16th century is considered as the “time of great herbalists”. The best known
among the herbalists belong to Germany. Among them were Otto Brunfels (1464–1534) known for
his herbal Herbarum Vivae Eicones, Jerome Bock (1489–1554) for his herbal Neu Kreuterbuch, and
Leonhart Fuchs (1501–1566) for his herbal De Historia Stirpium. All these herbalists are considered
as the “German Fathers of Botany” and their herbals exhibit some excellent illustrations and detailed
10 Plant Taxonomy
taxonomic descriptions of several available plants. However, they did not emphasize on any system
of classification of plants.
2.1.5 Taxonomy during Seventeenth Century

Andrea Caesalpino (1519–1603), an Italian, was the first scientist who worked for achieving a ratio-
nal scheme of classification of plants. Another name of repute among the taxonomists of the 17th
century is that of Gaspar Bauhin (1560–1624), a Swiss botanist. He compiled all the different kinds
of plants known to the science till then in a register called Pinax Theatri Botanici. His register,
which had an account of different names used by various workers for each plant, is considered
an authoritative discussion of synonymy in systematic botany. He, first, established the distinction
between the concept of genus and species and also initiated the use of the binomial nomenclature,
in some cases though not consistently.
John Ray (1627–1705), an English taxonomist, is another great contributor of the seventeenth cen-
tury. His works are published in two books entitled Methodus Plantarum Nova (1682) and Historia
Plantarum (1686). He also suggested a scheme of classification of plants.
Joseph Pitton de Tournefort (1656–1708) is another significant name in the field of taxonomy,
mainly for his publication titled Institutiones Rei Herbariae. He arranged over 9,000 kinds of plants
in about 700 genera grouping them in classes.
2.1.6 Period of Linnaeus

The eighteenth century belongs clearly to Carl Linnaeus (1707–1778) from the point of view of
history of taxonomy. He is the creator of the modern system of nomenclature.
A son of a Swedish clergyman, Linnaeus was educated at the universities of Lund and Uppasala,
and obtained the degree of M.D. in Netherlands. After practicing medicine for a few years, he became
a Professor of natural history at the University of Uppasala, where he spent the rest of his life.
Linnaeus is considered as the first taxonomist who showed that the reproductive features were
of paramount importance in taxonomy. He was the originator of the sexual system of classification,
in which he recognized 24 classes, mainly on the basis of number, length, union and certain other
characters of stamens. He was the first to use consistently the “binomial system of nomenclature”,
in which each organism is represented first by its generic name followed by the name of its species.
This scheme of nomenclature provided by Linnaeus is used throughout the world till today. The
plant taxonomy findings of Linnaeus were published in May 1753 in the form of his work Species
Plantarum.
The name of Linnaeus is commemorated today in the form of a well-established scientific soci-
ety, The Linnaean Society of London, and journals, such as Linnaea. Contribution of Linnaeus has
been so great that it may well be stated that so far the ‘plant taxonomy’ would survive in the world,
Linnaeus would be remembered.
2.1.7 Natural System Approach

Though Linnaeus (1753) was the first one to establish a system of classification of plants based on
reproductive parts, several totally unrelated plants were classified together (e.g. Prunus was classified
along with Cactus because of the same number of stamens) in his system of classification. This fact
compelled the botanists after Linnaeus to realise that no single character is intrinsically or naturally
more important than any other character. An approach to a natural system of classification first
took seed in France, where Michel Adanson (1727–1806) emphasised the fact that in several cases
natural characters are more useful than the others. This theory of Adanson was later recognised as
phenetic taxonomy.
The first scheme of classification based on natural characters was presented in 1789 by Antoine-
Laurent de Jussieu (1748–1836), a great botanist of France. All the four members of A.L. de ]ussieu’s
family (Antoine, Bernard, Joseph, and Antoine-Laurent) made notable contributions to the science
of plant taxonomy. The plants resembling each other in a set of characters were grouped together
in A.L. de Jussieu’s scheme of classification, and therefore, it was purely natural in its approach.
He presented his scheme of classification in his Genera Plantarum Secundum Ordines Naturales
Disposita.
Another family of botanists, contemporary to A.L. de Jussieu (1748–1836) was that of Augustin
Pyramus de Candolle (1778–1841). A.P. de Candolle presented a new classification of plants in his
book Theorie elementaire. He followed the approach of the natural system in his scheme and put all
alike plants together. An attempt to prepare an account of all available higher plants of the world
was undertaken by A.P. de Candolle in his Prodromus Systematis Naturalis Regni Vegetabilis in
1816, and he continued with this unfinished project till his death in 1841. The same project was
continued by his son, Alphonse (1806–1893) until 1873. After 1873, Alphonse and his son, Anne
Casimir de Candolle (1836–1918) published the details of this project in the form of monographs,
but this ‘great project’ could never be completed.
The latest, the best and a highly recognised natural system of classification was proposed by
George Bentham (1800–1884) and Joseph Dalton Hooker (1817–1911). They classified plants strictly
on the basis of a natural scheme. They made thorough observations of the material from the her-
baria, took very little help from the literature existing at that time, and presented their well-known
scheme of classification in their book titled Genera Plantarum. The Bentham and Hooker’s system
of classification is still supposed to be the best classification system, especially from the practical
laboratory point of view.
2.1.8 The Phylogenetic Approach

Publication of Darwin’s theory of organic evolution in 1859 in the form of his well-known book, On
the Origin of Species, provided the real base for the development of modern systematics. Scientists
started working on the line of thought that the life of plants as well as animals has continually
changed on the earth over a period of time. Botanists started working on the concept of evolution
regarding the development of a classification system of plants.
S. Endlicher (1805–1849) and A.W. Eichler (1839–1889), the two German botanists, were the first
to start working along this line of thought. They proposed the phylogenetic systems of the classifica-
tion of plants, and their schemes were later modified and developed by A. Engler (1844–1930) and
K. Prantl (1849–1893)1. Engler and Prantl’s monumental work was published in several volumes in
the form of Die Naturlichen Pflanzenfamilien between 1887 and 1915. Published in the 1915 volume
1
Details are mentioned in Chapter 3 (Classification).
12 Plant Taxonomy
of Pflanzenfamilien their scheme indicates that they started from simplest plants (e.g. Salicaceae
in case of dicotyledons because the members of this family possess simplest floral structures) and
ended with plants of complex floral structures (e.g. Compositae or Asteraceae in case of dicotyledons
because members of this family possess the height of floral complexity). Majority of the present day
botanical institutions and publications follow Engler and Prantl’s scheme of classification.
Another important work, contemporary to Engler and Prantl, was that of C.E. Bessey1 (1845–1915),
a Professor of botany at the University of Nebraska. Bessey grouped the flowering plants on the basis
of their evolutionary relationships. He categorised them on the basis of characters of primitiveness
and advanceness.l These guiding characters or principles were named as ‘dicta’ by Bessey. Richard
von Wettstein (1862–1931), an Austrian botanist, and Hans Hallier (1868–1938), a German botanist,
are two other plant taxonomists who suggested phylogenetic systems of classification.
John Hutchinson (1884–1972), a British botanist also suggested a widely recognized phylogenetic
system of plant classification which he published in two volumes of his well-known book The Families
of Flowering Plants. First published in 1926, the 3rd edition of this great book appeared in 1973,
shortly after his death on 2nd September 1972 at the age of 88. He classified the flowering plants
on the basis of 24 general principles.
2.1.9 Some Current Contributions

Four major systems of plant classification have been published during the last few decades, by
Armen Takhtajan (1910– ) of Leningrad (Russia), Arthur Cronquist (1919– ) of New York (USA),
Rolf Dahlgren (1919– ) of Copenhagen (Denmark), and Robert F. Thorne (1920– ) of Claremont
(USA).
Armen Takhtajan (1910– ) of Botanical Institute of Academy of Sciences, Leningrad, proposed a
system of classification of flowering plants in Russian in 1954. Its English version was first published
in 1969 in Flowering Plants: Origin and Dispersal. A new version of his classification1 is published
in Botanical Review of 1980. According to him the angiosperms are monophyletic and evolved from
some very ancient group of gymnosperms. He divided the angiosperms (Division: Magnoliophyta)
into two classes, i.e. Magnoliopsida (=Dicotyledons) and Liliopsida (= Monocotyledons).
Arthur Cronquist (1919– ) of New York Botanical Garden proposed a comprehensive system
of classification of angiosperms.2 First appeared in 1968 in The Evolution and Classification of
Flowering Plants, the latest version of Cronquist’s classification is published in 1981 in “An Integrated
System of Classification of Flowering Plants”.3 Cronquist considered the seed ferns (Pteridosperms)
as the probable ancestors of angiosperms. He divided angiosperms into Magnoliopsida and Liliopsida.
The Magnoliopsida, equivalent to Dicotyledons, consists of 64 orders, 318 families and about 1,65,000
species, whereas the Liliopsida consists of 19 orders, 65 families and about 50,000 species.
Rolf Dahlgren (1919– ) of Botanical Museum of University of Copenhagen proposed a new scheme 4
of classification of angiosperms. First published in 1975 in Botanische Notiser, the revised and
improved versions of his scheme appeared in 1980 in Botantcal Journal of Linnean Society, in 1981
1
Details are discussed in Chapter 3, Article No. 3.6.1.
2
Details are discussed in Chapter 3, Article No. 3.6.2.
3
Published by Columbia University Press, New York.
4
Details are mentioned in Chapter 3, Article No. 3.6.3.
in Phytochemistry and Angiosperm Phylogeny, and in 1983 in Nordiac Journal of Botany. He also
considered angiosperms to be monophyletic, and divided the class Magnoliopsida (= Angiospermae)
into two subclasses viz. Magnoliidae (= Dicotyledoneae) and Liliidae (= Monocotyledoneae).
Robert F. Thorne (1920– ) of Rancho Santa Ana Botanic Garden, Claremont, California, published
a phylogenetic system of classification1 of plants. First published in 1968 in Aliso, the revised and
enlarged outlines of Thorne’s scheme were published in 1981 in Phytochemistry and Phylogeny and
in 1983 in Nordiac Journal of Botany. Thorne has tried to establish the phylogenetic relationships
among the higher taxa of flowering plants. He divided the class Annonopsida (= Angiospermae)
into subclasses Annonidae (= Dicotyledoneae) and Liliidae (= Monocotyledoneae). Thome divided
Annonidae (= Dicotyledoneae) into 19 superorders, 41 orders, 56 suborders, 297 families, 350 sub-
families, 9,640 genera and 1,73,370 species, and Liliidae (= Monocotyledoneae) into 9 superorders,
12 orders, 17 suborders, 53 families, 102 subfamilies, 2,615 genera and 52,120 species.
2.2 TAXONOMY OUR CONTEMPORARY

The plant kingdom is very large and diversified. We are faced with an estimated total of about
3,00,000, or if fungi included, 4,50,000 plants. This large number includes many different kinds of
plants of a vast range in size and structure. This entire living world of plants forms the subject matter
of the plant taxonomy. The raw material of the taxonomic botany is the diversity of plant life in all
its aspects. One is, therefore, faced with exceedingly diverse taxonomical activities and techniques.
The algologist collecting algae in the arctic region, the plant collector in the tropical forest, the
botanist describing plants of a herbarium, the biochemist studying amino acids by chromatography,
the cytogeneticist studying breeding systems, the cytologist finding the chromosome number, the
electron microscopist going down to molecular details, the botanical artist depicting plants with his
pen, brush and ink, all contribute to the growth and development of plant taxonomy. Taxonomy is
the only branch of botany that involves the interests and abilities of so many practitioners. From a
plant collector to a computer programmer, from a biochemist to an electron microscopist, and from
a mycologist to a taxonomist, every one has room to show his talents in this branch. Thus, we can
say that taxonomy is truly our contemporary.
2.3 CHRONOLOGICAL DEVELOPMENT OF TAXONOMY IN INDIA

Indians of very old times had a satisfactory knowledge of plant taxonomy is evident from the studies
of the earliest Indian civilization as well as from the treatises such as Vrikshayurveda of Parasara,
Charaka Samhita, Sushrut Samhita and voluminous Indian classics such as Vedas, Ramayana,
Mahabharata, etc. However, a complete chronology of the development of plant taxonomy in India is
beyond the scope of this book. The major attempts of the botanical explorations of India during the
past few centuries and some of the important published works are undermentioned in Table 2.1.
2.4 SOME INDIAN JOURNALS RELATED TO TAXONOMY

1. Annals of the Royal Botanic Garden, Calcutta (now Kolkata)
2. Bulletin of the Botanical Society of Bengal
1
Details are mentioned in Chapter 3, Article No. 3.6.4.
14 Plant Taxonomy
3. Bulletin of the Botanical Survey of India

4. Indian Forest Records
5. Indian Journal of Forestry
6. Journal of Bombay Natural History Society
7. Journal of Economic and Taxonomic Botany
8. The Indian Forester
9. The Indian Journal of Agricultural Science
10. The Journal of the Asiatic Society of Bengal
11. The Records of the Botanical Survey of India
Table 2.1 Chronology of the major published taxonomic works of India
Year A.D. Researcher/Explorer Major published/botanical work
1565 Garcia d’Orta Published Os Colquios in Goa; described common

Indian medicinal plants.
1578 C. Acosta Published Tractado de las Drogas
1670 Heinrich van Rheede Published Hortus Malabaricus
1728–1785 John Gerard Koenig Formed society called “The United Brothers” in
order to promote and study Indian plants
1746–1793 Robert Kyd Founded Botanical Gardens of Calcutta
1751–1815 William Roxburgh Published Flora Indica and Plantae
Coromandelianus; prepared 2382 coloured
drawings of Indian plants; popularly called
Linnaeus of India
1817 Nathaniel Wallich Published Plantae Asiaticae Rariores
1824 J.F. Watson Published Flora of Kumaon
1833 J.F. Royle Flora of Kashmir
1839 J. Grahm Flora of Bombay
1840 J.W. Masters Flora of Calcutta
1844 W. Munro Flora of Agra
1857 J. Long Flora of Bengal
1859 W. Elliot Flora of Andhra
1859 T. Anderson Flora of Lucknow
1869 J.L. Stewart Flora of Punjab
1872–1897 J.D. Hooker Flora of British India published in the form of
seven volumes
1887 George King Started the publication of the journal The Annals
of the Royal Botanic Garden
1890 George King Established Botanical Survey of India
1901–1908 T. Cooke Flora of Presidency of Bombay
1902 H. Collett Flora Simlensis
(Contd.)
1903 D. Prain Bengal Plants

1903–1922 J.F. Duthie Flora of Upper Gangetic Plains and the Adjacent
Siwaliks and Sub-Himalayan Tract
1908 T.F. Bourdillon The Forest Trees of Travancore
1910 H.H. Haines A Forest Flora of Chota Nagpur
1915–1936 J.S. Gamble and Flora of Presidency of Madras
C.E.C. Fisher
1921 K. Rangacharier A Handbook of South Indian Grasses
1921–1925 H.H. Haines The Botany of Bihar and Orissa
1923 C.E. Parkinson Forest Flora of Andaman Islands
1929 P.V. Mayurnathan The Flowering Plants of Madras City and its
Immediate Neighbourhood
1929 E. Blatter and W.S. Millard Beautiful Indian Trees
1932 P.F. Fyson The Flora of South Indian Hill Stations
1932–1940 V. Kanjilal Flora of Assam
1940 N.L. Bor Common Grasses of United Provinces
1949 R.L. Bahadur and Poisonous Plants of India
S. Ghosh
1953 S.P. Agahakar Medicinal Plants
1960 B.P. Pal Beautiful Climbers of India
1962 H. Santapau The Flora of Saurastra
1962 K. Subramanian Aquatic Angiosperms
1964 P. Maheshwari and U. Singh Dictionary of Economic Plants in India
1964 G.S. Puri et al. Flora of Rajasthan
1965 M.S. Randhawa Flowering Trees
1965 J.K. Maheshwari Illustrated Flora of Delhi
1966 B.P. Pal The Roses in India
1966 V.D. Vartak Enumeration of Plants from Gomantak (Goa)
1968 R.K. Gupta Flora Nainitalensis
1968 R. N. Chopra and S.L. Nayyar Glossary of Indian Medicinal Plants
1968 S.L. Jindal Ornamental Bulbous Plants
1973 S.V. Ramaswamy and B.A. Raji Flora of Bangalore District
1973 Santapau and Henry Dictionary of Flowering Plants in India
1974 B.P. Pal and Vishnuswarup Bougainvilleas
1975 M.B. Raizada Supplement to the Duthie’s Flora of Upper
Gangetic Plains
1975 M.A. Rau High Altitude Flowering Plants of Western
Himalayas
1976 T.N. Srivastava Flora Gorakhporensis
1976 C.J. Saldanah and D.H. Nicolson Flora of Hasan District, Karnataka
1976 R. Rao Flowering Plants of Travancore
1977 M.M. Bhandari Flora of Rajasthan Desert
(Contd.)
16 Plant Taxonomy
1977 M. Oomachan The Flora of Bhopal

1978 M.B. Raizada and H.O. Saxena Flora of Mussoorie
1978 N.C. Nair Flora of Punjab Plains
1979 A.S. Rao Orchids of India
1979 V.N. Naik The Flora of Osmanabad
1979 B.L. Sapru et al. Flora of Ladakh
1983 U. Singh, A.M. Wadhwani and Dictionary of Economic Plants in India
B.M. Johri
1984 B.D. Naithani Flora of Chamoli
1984 H.J. Chowdhary and B.M. Wadhwa Flora of Himachal Pradesh
1984 O. Polunin and A. Stainton Flowers of Himalayas
1984 C.J. Saldanha Flora of Karnataka
1986 M.K. Kaul Weed Flora of Kashmir Valley
1986 CSIR Publication The Useful Plants of India
1987 Som Deva and H.B. Naithani The Orchid Flora of N.W. Himalayas
1988 R.K. Gupta The Living Himalaya Plant Exploration and
Phytogeography
1988 S.P. Vij (ed.) Biology, Conservation and Culture of Orchids
1992 H.K. Bakhu Herbs That Heal
1994 S.K. Jain Medicinal Plants (5th ed.)
1995 K. Haridarshan, G.P. Shukla and Medicinal Plants of Arunanchal Pradesh
B.S. Beniwal
1997 T.K. Chatterjee Herbal Options
1998 B. Bhattacharyya and B.M. Johri Flowering Plants: Taxonomy and Phylogeny
2001 S.K. Bhattacharjee Handbook of Medicinal Plants
2004 K.M. Nadkarni Medicinal Plants of India
2005 S.K. Bhattacharjee and L.C. Dey Medicinal Herbs and Flowers
2006 T.C. Narendran An Introduction to Taxonomy
2007 A.K. Singh Sedges and Grasses of Eastern Uttar Pradesh
2.5 A NOTE ON BOTANICAL SURVEY OF INDIA

George King, the Chief Executive Officer of the Royal Botanic Gardens, established the Botanical
Survey of India in 1890 with the main objective to coordinate the botanical work, being done in
different parts of India at that time. Regional offices of BSI were established at Madras1 with
M.A. Lawson as the Director for southern India, at Saharanpur with J.F. Duthie as the Director for
northern India, and at Pune with T. Cooke as the Director for the western India. The publication
of a journal The Records of the Botanical Survey of India was also started by George King. The
Industrial Section of the Indian Museum at Kolkata and the office of the Reporter of Economic
Products were brought under BSI in 1911. But the coming few decades were a period of inactivity
1
Renamed now Chennai.
for BSI mainly because of lack of funds and manpower. In 1939 the conditions deteriorated so that
the post of Director at Kolkata remained suspended, and BSI was surviving only in the form of a
Curator, Industrial Section, and a Systematic Assistant in the herbarium at Sibpur (Howrah).
The Government of independent India then came to the rescue of BSI. E.K. Janaki Ammal was
appointed as the Officer-on-Special Duty in 1952 to reorganise BSI. The first unit was established
in the form of The Central Botanical Laboratory at Allahabad with Janaki Ammal as the Director.
Four Regional Circles were established with their headquarters at Dehradun of Northern Circle,
Coimbatore of Southern Circle, Shillong of Eastern Circle, and at Pune of Western Circle. In 1962
the Central Botanical Laboratory was shifted from Allahabad to Kolkata, and at Allahabad a new
regional station for Central India was established. In 1972 two new circles (Andaman and Nicobar
Circle, and Arid Zone Circle) were established. Arunachal Pradesh Circle with its headquarter at
Itanagar was created in 1977, and Sikkim-Himalaya Circle with its head office at Gangtok in 1979.
The Deccan Circle at Hyderabad, and High Altitude Circle at Solan (Himachal Pradesh) were estab-
lished in 1984.
BSI headquarter office at Kolkata coordinates the research and other activities of all units, mainly
on the basis of the scientific policies of the Central Government. It also maintains links of BSI with
other major research institutions of the country such as CSIR, ICAR, ICMR, etc. A senior scientist
of BSI is also posted at the Royal Botanic Gardens, Kew (England) to maintain a link between the
two organizations as well as for many technical inquiries and clarifications regarding nomenclature,
etc.
Some of the major publications of BSI include the Bulletin of Botanical Survey of India, The
Records of the Botanical Survey of India, Annual Reports of BSI, and Newsletters. During the last
50 years, over 4000 research papers have been published by the scientists of BSI, and the herbaria
of the Survey hold over 2 million plant specimens.

1. Who is known as the “grandfather of modern Botany”?
2. The monumental work of which of the great physician of Roman army is compiled in the
form of Materia Medica?
3. Write a note on “herbalists” in about 100 words.
4. Give brief details of some discoveries made by Linnaeus.
5. Linnaeus obtained degree of M.D. and practiced ________ for few years and then became a
Professor of ________ at the University of ________ in Netherlands.
6. Armen Takhtajan, Arthur Cronquist, Rolf Dahlgren and Robert F. Thorne belonged to which
countries?
7. J.F. Duthie has been the author of the flora of which region of India?
8. Name three Indian journals related to plant taxonomy.
9. Write a detailed note on Botanical Survey of India.
18 Plant Taxonomy
Suggested Reading
Arber, A., 1938, Herbals: Their Origin and Evolution, Cambridge University Press, England.
Core, E.L., 1955, Plant Taxonomy, Prentice-Hall, Englewood Cliffs, N.J.
Cronquist, A., 1968, The Evolution and Classification of Flowering Plants, Thomas Nelson & Sons, Ltd.,
London.
Ewan, J.A., 1969, A Short History of Botany, Hafner Publishing Comp., New York.
Gibbs, R.D., 1963, History of Chemical Taxonomy, Academic Press, London.
Green, J.R., 1909, History of Botany 1860–1900, Clarendon Press, Oxford.
Maheshwari, P. and R.N. Kapil, 1963, Fifty Years of Science in India: Progress of Botany, Indian Sci.
Congr. Assoc., Calcutta.
Mayr, E., 1982, The Growth of Biological Thought, Harward Univ. Press, Cambridge.
Rao, R.S., 1973, Angiosperm Taxonomy, (In) A Decade (1963–72) of Science in India: Progress of Botany,
pp. 32–36. Indian Sci. Congr. Assoc. Calcutta.
Santapau, H., 1958, History of Botanical Research in India, Burma and Ceylon, Part II, Systematic Botany
of Angiosperms, The Bangalore Press, Bangalore.
Sneath, P.H.A., 1957, Application of Computers in Taxonomy, J. Gen. Microbiol. 17: 201–226.
Steere, W.C., 1958, Fifty Years of Botany, McGraw-Hill Book Co., New York.
Turrill, W.B., 1938, The Expansion of Taxonomy, Biol. Rev. 13: 342–373.
Wilmott, A.J., 1950, Systematic Botany from Linnaeus to Darwin, (In) Lectures on the Development of
Taxonomy, Linnean Society, London.
C
H
A
P
T
E
R
CLASSIFICATION 3
3.1 WHAT IS CLASSIFICATION?
Andrew Sugden (1984) defined the word “classification” in Longman Illustrated Dictionary of
Botany as “the naming of species and their grouping into families, orders, divisions, etc.” Radford
(1986) stated that “classification is the arrangement of groups of plants with particular circumscrip-
tions by rank and position according to artificial criteria, phenetic similarities, or phylogenetic
relationships.”
In its simplest form, classification is the placement of plants, animals and objects into groups and
categories for a clear understanding, proper study and effective organization.
3.2 RANKS OF PLANT CLASSIFICATION

Species, genus, family, order, class, and division are the six main ranks of plant classification in
an ascending order. Each rank has its subcategories, i.e. towards the higher ranks, subform, form,
subvarieties, varieties, and subspecies are the subcategories of species; subsection, section, and sub-
genus are the subcategories of genus; subtribe, tribe, and subfamily are the subcategories of family;
suborder is the subcategory of order; subclass is the subcategory of class; and subdivision is the
subcategory of division.
Several species are included within a genus, several genera within a family, several families within
an order, several orders within a class, and several classes are included within a division.
3.3 TYPES OF SYSTEMS OF CLASSIFICATION

The past taxonomic literature described three basic categories of systems of classification viz. artifi-
cial systems, natural systems and phylogenetic systems. But Radford (1986) has described following
four types of the systems of classification:
1. Artificial Classifications These systems use the habit and importance to man as the taxo-
nomic characters. Some advocates of artificial systems of classifications were Theophrastus
(370–285 B.C.), Secundus (23–79 A.D.), Dioscorides (62–128 A.D.), Magnus (1200–1280
20 Plant Taxonomy
A.D.), Brunfels (1464–1534), Bock (1489–1554) and Fuchs (1501–1566). Details of the works
of these workers are mentioned in Chapter 2 (Articles No. 2.1.2–2.1.4).
2. Mechanical Classifications These systems used one or a few selected taxonomic characters
to group taxa. Some mechanical classifications were given by Caesalpino (1519–1603), Bauhin
(1560–1624), Ray (1627–1705), Tournefort (1656–1708) and Linnaeus (1707–1778). Details of
the works of these workers are mentioned in Chapter 2 (Articles No. 2.1.5 and 2.1.6).
3. Natural Classifications These systems of classifications used as many taxonomic charac-
ters as possible to group taxa. Some of the natural systems of classification were given by
Adanson (1727–1806), A.L. de Jussieu (1748–1836) and his three family members (Antoine,
Bernard, and Joseph), A.P. de Candolle (1778–1841) and his son Alphonse (1806–1893), and
Bentham (1800–1884) and Hooker (1817–1911). Some details of the works of these workers
are mentioned in Chapter 2 (Article No. 2.1.7).
4. Phylogenetic Classifications These systems of classification used as many taxonomic
characters as possible in addition to the phylogenetic (evolutionary) interpretations. Some
of the phylogenetic systems of classification were proposed by Eichler (1839–1889), Engler
(1844–1930) and Prantl (1849–1893), Bessey (1845–1915), Wettstein (1862–1931), Hallier
(1868–1938), Hutchinson (1884–1972), Takhtajan (1980), Cronquist (1981), Dahlgren (1983)
and Thorne (1983). Some historical details of the works of these workers are mentioned in
Chapter 2 (Articles No. 2.1.8 and 2.1.9).
3.4 SOME IMPORTANT SYSTEMS OF CLASSIFICATION

3.4.1 John Ray (1627–1725)
An English biologist, John Ray1 treated over 18,000 species in the last edition of his book Methodus
Plantarum Nova published in 1703. He developed a classification system on the basis of the form
relationships by grouping together the plants that resembled one another. Ray was the first to divide
herbs, shrubs and trees into Dicotyledons and Monocotyledons on the basis of the presence of two
or one cotyledons. Broadly, he divided the plants as under:
Imperfectae (Flowerless plants)
I. Herbae (herbs) Dicotyledones
Perfectae (Flowering plants)

Monocotyledones
Monocotyledones
II. Arborae (shrubs and trees)
Dicotyledones
1
For historical details, see Chapter 2 (Article No. 2.1.5).
Classification 21
3.4.2 Carl Linnaeus1 (1707–1778)

Linnaeus,2 the father of taxonomy and creator of binomial system of nomenclature is considered as
“the most prodigious systematist of all times” (Lawrence, 1951). He was born on May 23, 1707 in
Rashult, southern Sweden.
In 1727, at the age of twenty, Linnaeus entered the University of Lund to study medicine. In 1728
he was transferred to the University of Uppasala, and published his first research paper on sexuality
of plants in 1729. In 1737 he received M.D. degree from the University of Harderwijk of Netherlands.
In Netherlands, he became the personal physician of a wealthy banker George Clifford. Clifford was
interested in plants. Linnaeus remained for three years in Netherlands, travelled extensively through
Europe, and collected plants. These three years were the most creative years of his life. In 1741, he
returned to Sweden, became the Professor of botany and medicine at the University of Uppasala,
and continued in the same position till his death on January 10, 1778.
Linnaeus was so popular as a teacher that his classes on field trips were attended often by 200 or
more students. He was buried in the Cathedral at Uppasala “unshaven, unwashed, unclad, enveloped
with a sheet” as per his own wish and request.
Three universally known works of Linnaeus are the Systema Naturae published in 1735, which
contains the outline of his system of classification; Genera Plantarum published in 1737 that contains
the description of several genera; and Species Plantarum published in 1753 in two volumes, which
contains his detailed views on plant identification and description of species.
Plant collections of Linnaeus were sold by his wife in 1783 to J.E. Smith, a British botanist, who
was one of the founders of Linnean Society of London. The entire plant collection is now housed
in the office of the Linnean Society in Burlington House, Piccadilly, London.
The Linnaeus’s system of classification is an artificial sexual system. However, Radford (1986)
described it as a mechanical system of classification. Linnaeus divided plants into 24 classes, mainly
on the bases of number, union, and length of stamens. The classes were divided into orders. On the
basis of the number of stamens in each flower Linnaeus placed all algae, fungi, mosses and ferns
under one class Cryptogamia.
All the 24 classes of the classification of Linnaeus are undermentioned:
1. Monandria (with 1 stamen), e.g. Scirpus.
2. Diandria (with 2 stamens), e.g. Veronica.
3. Triandria (with 3 stamens), e.g. Iris.
4. Tetrandria (with 4 stamens), e.g. Ulmus, Mentha.
5. Pentandria (with 5 stamens), e.g. Primula.
6. Hexandria (with 6 stamens), e.g. Berberis, Rumex.
7. Heptandria (with 7 stamens), e.g. Aesculus.
1
He was also known as Carl Linné or Carl von Linné. Lawrence (1951) and most of the other taxonomists mentioned
the spelling of his name as “Carolus Linnaeus”. But some recent taxonomists (Radford, 1986; Jones & Luchsinger, 1987)
spelled his name as Carl Linnaeus.
2
For more details of the history of Linnaeus, consult Chapter 2 (Article No. 2.1.6).
22 Plant Taxonomy
8. Octandria (with 8 stamens), e.g. Fagopyrum.

9. Enneandria (with 9 stamens), e.g. Ranunculus.
10. Decandria (with 10 stamens), e.g. Acer.
11. Dodecandria (with 11–19 stamens), e.g. Euphorbia.
12. Icasandria (with 20 or more stamens, episepalous), e.g. Rosa.
13. Polyandria (with 20 or more stamens, attached to axis), e.g. Papaver.
14. Didynamia (stamens didynamous), e.g. Linnaea.
15. Tetradynamia (stamens tetradynamous), e.g. Cruciferae.
16. Monadelphia (stamens monadelphous), e.g. Malvaceae.
17. Diadelphia (stamens diadelphous), e.g. Trifolia.
18. Polyadelphia (stamens polyadelphous), e.g. Hypericum.
19. Syngenesia (syngenesious condition), e.g. Compositae.
20. Gynandria (stamens adnate to gynoecium), e.g. Orchidaceae.
21. Monoecia (plants monoecious), e.g. Typha.
22. Dioecia (plants dioecious), e.g. Salix, Urtica.
23. Polygamia (plants polygamous), e.g. Empetrum.
24. Cryptogamia (flowers concealed), e.g. algae, fungi, mosses, ferns.
The classification system of Linnaeus dominated the botanical scene for over seven decades and
was replaced by natural classification systems of A.L. de ]ussieu and A.P. de Candolle based on
form relationships.
3.4.3 Antoine-Laurent de Jussieu (1748–1836)1

A natural system of classification was proposed by A.L. de Jussieu in 1789 in his Genera Plantarum
Secundus Ordines Naturales Disposita. He recognized one hundred orders of the plants which are
now called families. He divided the plants into three main groups, i.e. Acotyledones, Monocotyledones
and Dicotyledones. He mainly emphasized on the number of cotyledons and their presence or absence,
number of petals and their presence or absence, and position of stamens. This system firmly estab-
lished the philosophy of the natural system among the botanical community. All the hundred orders
(now called families) were arranged by A.L. de Jussieu into following 15 classes:
Groups Class
1. Acotyledones (plants without cotyledons, i.e. algae, fungi, mosses, etc.) ................................I
2. Monocotyledones (plants with one cotyledon)
Stamina hypogyna (stamens hypogynous) ........................................................................... II
Stamina perigyna (stamens perigynous).............................................................................. III
Stamina epigyna (stamens epigynous).................................................................................IV
3. Dicotyledones (plants with two cotyledons)
Stamina hypogyna (stamens hypogynous) V
1
For historical details see Chapter 2 (Article No. 2.1.7).
Classification 23
Stamina perigyna (stamens perigynous)..............................................................................VI

Stamina epigyna (stamens epigynous)............................................................................... VII
Monopetalae
Corolla hypogyna (corolla hypogynous)....................................................................... VIII
Corolla perigyna (corolla perigynous) .............................................................................IX
Corolla epigyna (corolla epigynous)
Antheris connatis (anthers connate) ................................................................................. X
Antheris distinctis (anthers free) .....................................................................................XI
Polypetalae
Stamina hypogyna (stamens hypogynous) .................................................................... XII
Stamina perigyna (stamens perigynous)....................................................................... XIII
Stamina epigyna (stamens epigynous)..........................................................................XIV
Declines irregularis (corolla generally absent; male and female flowers on different plants)
................................................................................................................................................ XV
3.4.4 Augustin Pyramus de Candolle (1778–1841)1

One more natural system of classification was proposed by A.P. de Candolle in 1813 in his book
Theorie Elementaire de la Botanique. He divided plants into two major groups i.e. Cellulares (non-
vascular plants), and Vasculares (vascular plants). In his another work, Prodromus, he described all
the species of the vascular plants known to the world then. In this work he described 58,000 species
of dicotyledons belonging to 161 families.
A brief outline of his plan is undermentioned:
1. Vasculares (plants with vascular bundles)
Class 1. Exogenae (Dicotyledoneae: vascular bundles in ring; 2 cotyledons)
(A) Diplochlamydeae (both calyx and corolla present)
(a) Thalamiflorae (polypetalous, hypogynous)—Orders 1–46.
(b) Calyciflorae (perigynous or epigynous)—Orders 47–84.
(c) Corolliflorae (gamopetalous and hypogynous)—Orders 85–108.
(B) Monochlamydeae (only calyx present)—Orders 109–128.
Class 2. Endogenae (Monocotyledoneae; vascular bundles scattered; cotyledon one)
(A) Phanerogamae (flowers present)—Orders 129–150.
(B) Cryptogamae (flowers absent or hidden)—Orders 151–155.
2. Cellulares (Plants without vascular bundles or cotyledons)
Class 1. Foliaceae (leafy and sexual)—Orders 156–157.
Class 2. Aphyllae (nonleafy and without known sexes)—Orders 158–161.
1
For historical details see Chapter 2 (Article No. 2.1.7).
24 Plant Taxonomy
The system of A.P. de Candolle was easy and simple, and surpassed all other systems. But its
major drawback was the inclusion of vascular cryptogams among the monocots.
3.4.5 George Bentham (1800–1884) and Joseph Dalton Hooker (1817–1911)

The most accepted natural system of classification was proposed by Bentham and Hooker in their
Genera Plantarum published during July 1862 and April 1883. Bentham, a self-trained British bota-
nist, and Hooker, the Director of the Royal Botanic Gardens, Kew (England), described all known
genera of seed plants in three volumes of their Genera Plantarum, published in Latin.
Bentham and Hooker’s system of classification is still used and followed in several herbaria of the
world. In most of the Indian herbaria too, the plants are arranged according to this system of clas-
sification. It is supposed to be the best system for the students to identify plants in the laboratories.
This is so because Bentham and Hooker prepared the generic descriptions of the plants from their
own observations and not by copying from the available literature. Large genera are divided into
sections and sub-sections, and the description of genera is complete and accurate.
In all, they described 97,205 species belonging to 7,569 genera of 200 families of flowering plants
in three volumes of Genera Plantarum as shown in Table 3.1.
Table 3.1 Number of orders, genera and families described by Bentham and Hooker
Groups Orders Genera Species

(families)
Dicotyledons
(a) Polypetalae 82 2,610 31,874
(b) Gamopetalae 45 2,619 34,556
(c) Monochlamydeae 36 801 11,784
Gymnosperms 3 44 415
Monocotyledons 34 1,495 18,576
Total 200 7,569 97,205
The Bentham and Hooker’s system of classification is clearly derived from the systems of de Jussieu
and de Candolle. Bentham and Hooker divided all Phanerogams or seed plants into Dicotyledons,
Gymnosperms and Monocotyledons. Ranales were placed in the beginning and grasses at the end
in this classification. A summary outline of their classification is mentioned below:
(A) Dicotyledons (Reticulate venation; two cotyledons; pentamerous flowers).
1. Polypetalae (Corolla of separate petals)
Series I. Thalamiflorae (stamens many; hypogynous; disc absent).
Order 1. Ranales: Ranunculaceae, Magnoliaceae, Annonaceae, Nymphaeaceae and four
more families.
2. Parietales: Papaveraceae, Capparidaceae, Cruciferae, Violaceae and five more
families.
3. Polygalineae: Polygaleae and three more families.
Classification 25
Caryophyllineae: Caryophyllaceae, Portulacaceae and two more families.

4.
Guttiferales: Guttiferae and five more families.
5.
Malvales: Malvaceae, Tiliaceae and Sterculiaceae.
6.
Series II.
Disciflorae (stamens hypogynous; disc present).
Order 1. Geraniales: Geraniaceae, Rutaceae, Meliaceae, and eight more families.
2. Olacales: Olacineae, and two more families.
3. Celastrales: Rhamnaceae, and three more families.
4. Sapindales: Sapindaceae, Anacardiaceae and Sabiaceae.
Series III. Calyciflorae (stamens perigynous or epigynous; ovary generally inferior).
Order 1. Rosales: Leguminosae, Rosaceae, and seven more families.
2. Myrtales: Combretaceae, Myrtaceae, Lythraceae, and three more families.
3. Passiflorales: Cucurbitaceae, Begoniaceae, and five more families.
4. Ficoidales: Cactaceae, Ficoideae.
5. Umbellales: Umbelliferae, and two more families.
2. Gamopetalae (petals of corolla are partially or completely fused).
Series I. Inferae (inferior ovary).
Order 1. Rubiales: Rubiaceae and Caprifoliaceae.
2. Asterales: Compositae, and three more families.
3. Campanales: Campanulaceae, and three more families.
Series II. Heteromerae (ovary superior; androecium of one or two series; carpels more
than two).
Order 1. Ericales: Ericaceae, and five more families.
2. Primulales: Primulaceae, and two more families.
3. Ebenales: Sapotaceae, and two more families.
Series III. Bicarpellatae (ovary superior; androecium of one series; carpels two).
Order 1. Gentianales: Oleaceae, Apocynaceae, Asclepiadaceae, and three more
families.
2. Polemoniales: Convolvulaceae, Solanaceae, and three more families.
3. Personales: Scrophulariaceae, Pedaliaceae, Bignoniaceae, Acanthaceae,
and four more families.
4. Lamiales: Labiatae, Verbenaceae, and two more families.
3. Monochlamydeae (Petals absent).
Series I. Curvembryeae (embryo coiled, ovule generally one): Amaranthaceae,
Chenopodiaceae, Polygonaceae, and four more families.
Series II. Multiovulatae aquaticae (ovules many; immersed aquatics): Podostemaceae.
Series III. Multiovulatae terrestris (ovules many; plants terrestrial): Nepenthaceae, and
two more families.
Series IV. Microembryeae (embryo very minute): Piperaceae, and three more families.
26 Plant Taxonomy
Series V. Daphnales (ovary with one carpel and one ovule): Proteaceae, and three more
families.
Series VI. Achlamydosporeae (usually inferior ovary; one locule, with 1–3 ovules):
Loranthaceae, Santalaceae and Balanophoreae.
Series VII. Unisexuales (flowers unisexual): Euphorbiaceae, Urticaceae, and seven more
families.
Series VIII. Ordines anomali (families of uncertain relationship): Ceratophyllaceae, and
three more families.
(B) Gymnospermae (naked-seeded plants): Gnetaceae, Coniferae, Cycadaceae.
(C) Monocotyledons (parallel venation; one cotyledon; trimerous flowers).
Series I. Microspermae (inferior ovary; minute seeds): Orchidaceae, and two more
families.
Series II. Epigynae (inferior ovary; large seeds): Iridaceae, Amaryllidaceae, and five
more families.
Series III. Coronarieae (superior ovary; coloured perianth): Liliaceae, Commelinaceae,
and six more families.
Series IV. Calycineae (superior ovary; green perianth): Juncaceae, Palmae,
Flagellariaceae.
Series V. Nudiflorae (perianth usually absent; superior ovary): Typhaceae, Araceae, and
three more families.
Series VI. Apocarpae (carpels free): Alismaceae, and two more families.
Series VII. Glumaceae (reduced perianth; bracts large, scaly): Cyperaceae, Gramineae,
and three more families.
Merits of the System of Bentham and Hooker
1. It is the first great natural system of classification.
2. It is very easy to follow for all practical purposes, and that is why Kew Herbarium and
several other herbaria of the world, including India, are arranged according to this system.
3. This system was never planned by Bentham and Hooker on the basis of phylogeny, although
the theory of organic evolution was already announced by Darwin and Wallace in 1859. So
this system should not be criticized on the basis of phylogeny.
4. Ranales have been given a primitive position in this system. Recent taxonomic findings also
indicate that Ranales are the most primitive living angiosperms.
5. In this system the monocots are derived from dicots. Several recent taxonomic findings sup-
port this view.
Demerits of the System of Bentham and Hooker
1. The position of gymnosperms in between dicots and monocots in this system is its foremost
demerit. This arrangement is made without considering the affinities among these groups.
2. Several important floral characters have been neglected in this system.
Classification 27
3. In this system some of the closely related families have been separated and placed under dif-
ferent orders (cohorts). In the same way, a number of unrelated families have been grouped
nearer. Few examples are undermentioned:
(i) All the families of Series Curvembrae are related to Caryophyllaceae of series
Thalamiflorae of Polypetalae.
(ii) Podostemaceae of Series Multiovulatae aquaticae of Monochlamydeae deserves a placing
under Rosales of Series Calyciflorae of Polypetalae.
(iii) Nepenthaceae (of Series Multiovulatae-terrestris of Monochlamydeae) is related more to
family Saracenniaceae (of orders Parietales of Thalamiflorae).
(iv) Laurineae of Series Daphnales is related closely with Magnoliaceae of Ranales.
4. Advanced families, such as Orchidaceae, have been considered primitive in this system by
placing them in the beginning.
5. The entire arrangement of monocots is unnatural and unphylogenetic in this system.
3.4.6 August Wilhelm Eichler (1839–1889)

Eichler (1883), a German botanist, put forth a widely accepted system of classification of plants.
This system may be considered as a transitional phylogenetic system. The schematic representation
of Eichler’s classification is undermentioned:
Plant Kingdom
1. Cryptogamae (Nonseed plants)
Divisions 1. Thallophyta: Algae, Fungi, Lichens
2. Bryophyta: Liverworts and Mosses
3. Pteridophyta: Equisetineae, Lycopodineae and Filicineae
2. Phanerogamae (Seed plants)
Divisions 1. Gymnospermae
2. Angiospermae
Class 1. Monocotyledones (7 orders)
2. Dicotyledones
Subclass 1. Choripetalae (20 orders)
2. Sympetalae (9 orders)
3.4.7 Adolf Engler (1844–1930) and Karl Prantl (1849–1893)

Engler and Prantl’s system of classification has also been described as a “transitional phylogenetic
system” by Jones and Luchsinger (1987). However, Radford (1986) and most of the other taxonomists
describe it as a phylogenetic system. Engler, a Professor at the University of Berlin, proposed a sys-
tem of classification based on that of Eichler. Between 1887 to 1915, he published it along with his
associate, Prantl, in several volumes of Die Naturlichen Pflanzenfamilien. This, profusely-illustrated
work of Engler and Prantl, provides keys and description of all the plant families known to them at
28 Plant Taxonomy
that time. They classified all the plants from algae to angiosperms. This system is used in most of
the non-British herbaria of the world.
The followers of Engler and Prantl published revised classification in several successive editions
of Syllabus der Pflanzenfamilien. The twelfth edition of Syllabus, dealing angiosperms, was edited
by Melchior in 1964.
The most noteworthy features of the Engler and Prantl’s system of classification are that they
(i) placed monocots before dicots, (ii) considered orchids to be more evolved than grasses, and
(iii) considered apetalous and catkin-bearing dicots primitive to the dicots bearing petals and simple
unisexual flowers.
Engler and Prantl’s system divided the plant kingdom into following 14 divisions:
1. Schizophyta (classes Schizomycetes and Schizophyceae), 2. Myxothallophyta (class Myxomycetes), 3.
Flagellatae, 4. Dinoflagellatae, 5. Bacillariophyta, 6. Conjugatae, 7. Heterocontae. 8. Chlorophyceae,
9. Charophyta, 10. Phaeophyceae, 11. Rhodophyceae, 12. Eumycetes (Fungi), 13. Archegoniatae
or Embryophyta-Asiphonogama (subdivisions Bryophyta and Pteridophyta), 14. Embryophyta-
Siphonogama (subdivisions Gymnospermae and Angiospermae).
Subdivision—Angiospermae
Class 1. Monocotyledoneae
Orders: 1. Pandanales (Typhaceae, etc.)
2. Helobiae (Alismataceae, and 6 more families)
3. Triuridales (Triuridaceae)
4. Glumiflorae (Cyperaceae, Gramineae)
5. Principes (Palmae)
6. Synanthae (Cyclanthaceae)
7. Spathiflorae (Araceae, Lemnaceae)
8. Farinosae (Commelinaceae, and 12 more families).
9. Liliiflorae (Juncaceae, Liliaceae, Amaryllidaceae, Iridaceae, and 5 more
families)
10. Scitamineae (Musaceae, and 3 more families)
11. Microspermae (Orchidaceae and Burmanniaceae)
Class 2. Dicotyledoneae
Subclass 1. Archichlamydeae
Orders: 1. Verticillatae (Casuarinaceae)
2. Piperales (Piperaceae, and 2 more families)
3. Hydrostachyales (Hydrostachyaceae)
4. Salicales (Salicaceae)
5. Garryales (Garryaceae)
6. Myricales (Myricaceae)
7. Balanopsidales (Balanopsidaceae)
Classification 29
8.
Leitneriales (Leitneriaceae)
9.
Juglandales (Juglandaceae)
10.
Julianiales (Julianiaceae)
11.
Batidales (Batidaceae)
Fagales (Fagaceae, Butolaceae)
12.
Urticales (Moraceae, Urticaceae, Ulmaceae)
13.
14.
Podostemonales (Podostemonaceae)
15.
Proteales (Proteaceae)
Santanales (Santalaceae, Loranthaceae, and 5 more families)
16.
17.
Aristolochiales (Aristolochiaceae, and 2 more families)
18.
Balanophorales (Balanophoraceae)
19.
Polygonales (Polygonaceae)
Centrospermae (Chenopodiaceae, Amaranthaceae, Nyctaginaceae, Caryophyl-
20.
laceae, and 6 more families)
21. Ranales (Ranunculaceae, Nymphaeaceae, Magnoliaceae, Annonaceae, and 15
more families)
22. Rhoeadales (Papaveraceae, Capparidaceae, Cruciferae, and 4 more
families)
23. Sarraceniales (3 families)
24. Rosales (Rosaceae, Leguminosae, and 15 more families)
25. Pandanales (Pandanaceae)
26. Geraniales (Geraniaceae, Rutaceae, Meliaceae, Euphorbiaceae, and 17 more
families)
27. Sapindales (Anacardiaceae, and 22 more families)
28. Rhamnales (Rhamnaceae, Vitaceae)
29. Malvales (Malvaceae, Tiliaceae, Bombacaceae, Sterculiaceae, and 3 more
families)
30. Parietales (Violaceae, and 30 more families)
31. Opuntiales (Cactaceae)
32. Myrtiflorae (Myrtaceae, Combretaceae, and 21 more families)
33. Umbelliflorae (Umbelliferae, and 2 more families)
Subclass 2. Metachlamydeae (Sympetalae)
Orders 1. Diapensiales (Diapensiaceae)
2. Ericales (Ericaceae, and 3 more families)
3. Primulales (Primulaceae, and 2 more families)
4. Plumbaginales (Plumbaginaceae)
5. Ebenales (Sapotaceae, and 6 more families)
6. Contortae (Apocynaceae, Asclepiadaceae, Oleaceae, and 3 more families)
30 Plant Taxonomy
7. Tubiflorae (Convolvulaceae, Boraginaceae, Verbenaceae, Labiatae, Solan-

aceae, Scrophulariaceae, Bignoniaceae, Pedaliaceae, Acanthaceae, and 13
more families)
8. Plantaginales (Plantaginaceae)
9. Rubiales (Rubiaceae, and 4 more families)
10. Cucurbitales (Cucurbitaceae)
11. Campanulatae (Campanulaceae, Compositae, and 4 more families)
Merits of the System of Engler and Prantl
1. This is a convenient and well-known filing system of several herbaria of the world.
2. Polypetalae and Monochlamydae of Bentham and Hooker were merged by Engler and Prantl
into one single subclass Archichlamydae.
3. This system treated families such as Orchidaceae and Compositae as advanced families.
4. In this system several closely related families (e.g. Liliaceae, Juncaceae, Iridaceae, and
Amaryllidaceae) are treated close to one another.
5. Abundant illustrations are provided along with the description of the families.
6. The system is provided with exhaustive keys of families and orders.
7. The description of each family also contains a summary of its embryology, morphology,
anatomy, and geographical distribution.
Demerits of the System of Engler and Prantl
1. Monocots have been placed before dicots in this system. But this system has been reversed
in the 1964 edition of Syllabus der Pflanzenfamilien.
2. Naked flowers of Amentiferae have been treated as primitive in this system.
3. Helobiae, consisting of primitive forms, have been placed between two advanced orders
Glumiflorae and Pandanales.
4. Araceae are derived from Liliaceae, but Engler and Prantl placed Araceae before
Liliaceae.
5. This system fails to recognize the significance of reduction, and because of this “simple”
were equated with “primitive” according to Cronquist (1965).
3.4.8 Charles Edwin Bessey (1845–1915)

A system of classification that tries to reflect evolution is called phylogenetic system. Bessey, a student
of Asa Gray, and a Professor at the University of Nebraska, contributed a pure phylogenetic system
of plant classification in 1894 after being highly impressed by the evolutionary ideas of Darwin.
It could be published in the final form in 1915, shortly after his death. Bessey closely followed
Bentham and Hooker’s system of classification. His system was based on certain guiding principles
of primitive and advanced characters called dicta1. He considered angiosperms monophyletic and
derived from a cycadeoid ancestor having bisexual strobili.
1
Some of the Besseyan principles or “dicta” are mentioned in Chapter 13 (Article 13.3).
Classification 31
According to Bessey (1915), Ranales were the primitive angiosperms. One branch of Ranales
developed into monocots, and the other into dicots. This system is often called Ranalian concept of
evolution. On being diagramed, this system took the form of a cactus plant, and came to be called
Bessey’s cactus (Fig. 3.1).
24
23
22 1. Ranales 17. Cactales

13 14
2. Serraceniales 18. Lossales
21 3. Malvales 19. Celastrales
20
4. Geraniales 20. Umbellales
11 12 5. Guttiferales 21. Sapindales
18 6. Rhoeadales 22. Rubiales
19
17 7. Caryophyllales 23. Campanulales
32 9
16 10 8. Ebenales 24. Asterales
9. Primulales 25. Alismatales
8 7
10. Ericales 26. Liliales
11. Geraniales 27. Hydrales
15 6
31 5 12. Polemoniales 28. Arales
13. Scrophulariales 29. Palmales
27 14. Lamiales 30. Graminales
30 4
1 15. Rosales 31. Iridales
29 26 3 16. Myrtales 32. Orchidales
28 25 2
Fig. 3.1 Bessey’s cactus—Ranalian concept of evolution.
In this system of classification, the angiosperms are divided as under:

Class—ALTERNIFOLIAE (Monocotyledoneae)
Subclass—Strobiloideae
Orders—Alismatales (Families 1–9), Liliales (Fam. 10–22), Arales (Fam. 23–25), Pal-
males (Fam. 26), Graminales (Fam. 27–31).
Subclass—Cotyloideae
Orders—Hydrales (Fam. 32), Iridales (Fam. 33–43), Orchidales (Fam. 44–45).
Class—OPPOSITIFOLIAE (Dicotyledoneae)
Subclass—Strobiloideae
Superorder—Apopetalae—Polycarpellatae
Orders—Ranales (Fam. 46–69). Malvales (Fam. 70–81), Sarraceniales (Fam. 82–83),
Geraniales (Fam. 84–105), Guttiferales (Fam. 106–125), Rhoeadales (Fam.
126–132), Caryophyllales (Fam. 133–149).
32 Plant Taxonomy
Superorder—Sympetalae—Polycarpellatae
Orders—Ebenales (Fam. 150–154), Ericales (Fam. 155–160), Primulales (Fam.
161–165).
Superorder—Sympetalae—Dicarpellatae
Orders—Gentianales (Fam. 166–171), Polemoniales (Fam. 172–177), Scrophulariales
(Fam. 178–187), Lamiales (188–191).
Subclass—Cotyloideae
Superorder—Apopetalae
Orders—Rosales (Fam. 192–214), Myrtales (Fam. 215–229), Loasales (Fam. 230–234),
Cactales (Fam. 235), Celastrales (Fam. 236–259), Sapindales (Fam. 260–274),
Umbellales (Fam. 275–277).
Superorder—Sympetalae
Order—Rubiales (Fam. 278–282), Campanulales (Fam. 283–286), Asterales (Fam.
287–300).
Merits of the System of Bessey
1. Bentham and Hooker placed Gymnosperms in between Dicotyledons and Monocotyledons.
Such is not the case in Bessey’s system of classification.
2. Monochlamydeae has been completely abolished by Bessey. Families of Monochlamydeae
have been distributed near their allies in Dicotyledons (Oppositifoliae).
3. Ranales are the starting point among Dicotyledons.
4. In Bessey’s system the families with an inferior ovary follow the families possessing a supe-
rior ovary.
Demerits of the System of Bessey
1. Monocots have been assigned a position prior to dicots, which is not proper.
2. Hypogyny, perigyny and epigyny have been emphasized too much in this system.
Comparison of Englerian and Besseyan Concepts
1. Englerian school believed the primitive flowers to be apetalous and unisexual, while Besseyan
school believed them polypetalous and bisexual.
2. Englerian school believed the wind pollination primitive while it was the insect pollination
according to Besseyan school.
3. According to Englerian concept, dicots began with Amentiferae while according to Besseyan
concept they began with Ranales.
4. Englerian school believed that monocots were derived from a gymnospermous stock while
according to Besseyan school they derived from some primitive dicots.
5. The main philosophy of Englerian school is that simple flowers are primitive while Besseyan
school believed that flowers with complex structures are more so.
Classification 33
3.4.9 John Hutchinson (1884–1972)

An intentional phylogenetic system was produced by John Hutchinson, a British botanist, associated
with the Royal Botanic Gardens, Kew. His system is close to that of Bessey, and is published in his
Genera of Flowering Plants (1964–1967) and Families of Flowering Plants (1973).
Hutchinson based his system of classification on the twenty four principles mentioned below in
a condensed form:
1. Evolution is both downwards and upwards.
2. Evolution does not necessarily involve all organs at the same time.
3. Generally, the evolution has been consistent.
4. Broadly, shrubs and trees are more primitive than herbs in any one genus or family.
5. In comparison with climbers, shrubs and trees are older in any one genus or family.
6. Perennials are older than annuals and biennials.
7. Aquatic flowering plants are derived from terrestrial ancestors.
8. Dicots are primitive compared to monocots.
9. Spiral arrangement of vegetative and floral leaves is primitive to cyclic arrangement.
10. Usually, simple leaves are more primitive than compound leaves.
11. Bisexual flowers are less advanced than unisexual, and dioecious plants are more recent than
monoecious.
12. Inflorescence is more advanced than the solitary flowers.
13. Types of aestivation are evolved from contorted to imbricate to valvate.
14. Polymerous flowers (many-parted) precede oligomerous (few-parted) flowers.
15. Apetalous flowers are derived from flowers with petals.
16. Polypetaly is more primitive than gamopetaly.
17. In comparison to zygomorphy, actinomorphy is more primitive.
18. Hypogyny is more primitive than perigyny. The most advanced condition is epigyny.
19. Apocarpy is primitive than syncarpy.
20. Polycarpellary condition precede the condition of a few carpels.
21. Endospermic seeds with a small embryo are primitive to non-endospermic ones with a large
embryo.
22. Flowers with many stamens are primitive to flowers with few stamens.
23. Plants with separate anthers are primitive compared to those with fused anthers or
filaments.
24. Aggregate fruits are more highly evolved than single fruits.
An outline of the Hutchinson’s system of classification, up to the level of orders, is mentioned
below. Common families of some of the orders are mentioned in parenthesis.
34 Plant Taxonomy
Phylum ANGIOSPERMAE
Subphylum 1. DICOTYLEDONES
Division I. Lignosae
Orders: 1. Magnoliales (Magnoliaceae); 2. Annonales (Annonaceae); 3. Laurales; 4. Dil-
leniales; 5. Coriariales; 6. Rosales (Rosaceae); 7. Leguminales; 8. Cunoniales; 9.
Styracales; 10. Araliales; 11. Hamamelidales; 12. Salicales; 13. Leitneriales; 14.
Myricales; 15. Balanopsidales; 16. Fagales (Fagaceae); 17. Juglandales; 18. Ca-
suarinales (Casuarinaceae); 19. Urticales (Moraceae, Urticaceae); 20. Bixales;
21. Thymeleales (Nyctaginaceae); 22. Proteales (Proteaceae); 23. Pittosporales;
24. Capparales (Capparidaceae); 25. Tamaricales; 26. Violales (Violaceae); 27.
Polygalales; 28. Loasales; 29. Passiflorales; 30. Cucurbitales (Cucurbitaceae,
Begoniaceae); 31. Cactales (Cactaceae); 32. Tiliales (Tiliaceae, Sterculiaceae,
Bombacaceae); 33. Malvales (Malvaceae); 34. Malpighiales; 35. Euphorbiales
(Euphorbiaceae); 36. Theales; 37. Ochnales; 38. Ericales; 39. Guttiferales; 40.
Myrtales (Myrtaceae); 41. Celastrales (Salvadoraceae); 42. Olacales; 43. San-
talales (Loranthaceae, Santalaceae); 44. Rhamnales (Vitaceae); 45. Myrsinales;
46. Ebenales (Sapotaceae); 47. Rutales (Rutaceae); 48. Meliales (Meliaceae);
49. Sapindales; 50. Loganiales (Oleaceae); 51. Apocynales (Apocynaceae,
Asclepiadaceae); 52. Rubiales (Rubiaceae); 53. Bignoniales (Bignoniaceae,
Pedaliaceae); 54. Verbenales (Verbenaceae).
Division II. Herbaceae
Orders: 55. Ranales (Ranunculaceae); 56. Berberidales; 57. Aristolochiales (Aris-
tolochiaceae); 58. Piperales (Piperaceae); 59. Rhoeadales (Papaveraceae,
Fumariaceae); 60. Brassicales (Brassicaceae); 61. Resedales; 62. Caryophyllales
(Caryophyllaceae); 63. Polygonales (Polygonaceae); 64. Chenopodiales (Che-
nopodiaceae, Amaranthaceae); 65. Onagrales; 66. Gentianales; 67. Primulales;
68. Plantaginales; 69. Saxifragales; 70. Sarraceniales; 71. Podostemales; 72. Um-
bellales (Umbelliferae); 73. Valerianales; 74. Campanulales; 75. Goodeniales;
76. Asterales (Compositae); 77. Solanales (Solanaceae, Convolvulaceae);
78. Personales (Acanthaceae, Scrophulariaceae); 79. Geraniales (Geraniaceae);
80. Polemoniales; 81. Boraginales (Boraginaceae); 82. Lamiales (Labiatae).
Subphylum 2. MONOCOTYLEDONES
Division. 1. Calyciferae
Orders: 83. Butomales (Butomaceae); 84. Alismatales; 85. Triuridales; 86. Juncaginales;
87. Aponogetonales; 88. Potamogetonales; 89. Najadales; 90. Commelinales
(Commelinaceae); 91. Xyridales; 92. Eriocaulales; 93. Bromeliales; 94. Zin-
giberales (Musaceae, Zingiberaceae).
Division. 2. Corolliferae
Orders: 95. Liliales (Liliaceae); 96. Alstromeriales; 97. Arales (Araceae); 98.
Typhales (Typhaceae); 99. Amaryllidales (Amaryllidaceae); 100. Tridales;
Classification 35
101. Dioscoreales; 102. Agavales; 103. Palmales (Palmae); 104. Pandanales;

105. Cyclanthales; 106. Haemodorales; 107. Burmanniales; 108. Orchidales
(Orchidaceae).
Division 3. Glumiflorae
Orders: 109. Juncales (Juncaceae); 110. Cyperales (Cyperaceae); 111. Graminales
(Gramineae).
Merits of the System of Hutchinson
1. Families and orders in this system are of small size and comprise only very closely related
taxa.
2. This system is in confirmity with the modern views of the phylogeny of angiosperms as it
considers the Ranales and Magnoliales as the starting points among dicots.
3. Monocots are discussed after dicots in this system.
Demerits of the System of Hutchinson
1. From the point of view of plant identification this system is not of much utility.
2. Dicots have been divided into two major groups (Lignosae and Herbaceae) in this system. This
kind of classification is outdated as the habit used to be the main basis of classification in
the past era of Aristotle. Secondly, it also separates the families that have close affinities.
3.5 COMPARISON OF SYSTEMS OF CLASSIFICATION

Table 3.2 provides points of comparison between three main systems of classification proposed by
Bentham and Hooker, Engler and Prantl, and Hutchinson.
3.6 CURRENT SYSTEMS OF CLASSIFICATION

Armen Takhtajan (1980) of Russia (formerly U.S.S.R.), Arthur Cronquist (1981) of U.S.A., Rolf
Dahlgren (1983) of Denmark, and Robert F. Thorne (1983) of U.S.A. published systems1 of plant
classification during last few decades. These workers used the recent data and information of palaeo-
botany, biochemical systematics, and the ultrastructural details unveiled by the electron microscope,
along with the information of the traditional sources such as morphology and anatomy of plants.
3.6.1 Armen Takhtajan (1980)1

The latest version of Takhtajan’s classification is published in 1980 in Botanical Review. According
to him, the angiosperms are monophyletic and developed from some very ancient group of gymno-
sperms. He is an adherent of the Besseyan philosophy of taxonomy.
Some of the criteria, used by Takhtajan in determining the relative degree of advancement
of flowering plants are undermentioned: (i) Woody plants are primitive compared to herbaceous
plants; (ii) Deciduous woody plants are evolved from evergreen plants; (iii) Parallel venation is most
advanced; (iv) Most primitive leaf arrangement is alternate; (v) Stomata with subsidiary cells are
1
For the historical details of these systems, refer Chapter 2 (Article No 2.1.9).
36 Plant Taxonomy
Table 3.2 Comparison of systems of classification
S. No. Bentham and Hooker Engler and Prantl Hutchinson
1. It is a natural system of It is a transitional phylogenetic It is a phylogenetic system.

classification. system.
2. It is based on de Candolle’s system It is based on Eichler’s system of It is based on Bessey’s system of
of classification. classification. classification.
3. Fixity of species is the main idea Darwinian theory of descent is the Considerable knowledge of
behind this system. main idea behind this system. phylogeny is the basis of this
classification.
4. Dicots ar divided into Dicots are divided into Dicots are divided into
(i) Polypetalae, (ii) Gamopetalae, (i) Archichlamydae, and (i) Lignosae, and (ii) Herbaceae
and (iii) Monochlamydae. (ii) Metachlamydae. in this system.
5. Gymnosperms are placed in Gymnosperms are placed before Gymnosperms are placed before
between dicots and monocots. dicots. dicots.
6. Monocots are treated after dicots. Monocots are treated before Monocots are treated after
dicots. dicots.
7. Orchidaceae and some other Orchidaceae are treated as Orchidaceae are treated as
advanced families are regarded advanced. advanced.
as primitive, and treated in the
beginning.
8. Families with free petals are placed Polypetalae and Monochlamydeae Emphasis is on resemblances
in Polypetalae, with fused petals in are placed together under a single in place of differences, and
Gamopetalae, and without petals group Archichlamydeae. Families so the Gamopetalae and
in Monochlamydeae. with fused petals are placed under Monochlamydeae are distributed
Metachlamydeae. according to their relationship
in Polypetalae.
9. Monocotyledonous families are Monocotyledons are arranged in 11 Monocotyledons are arranged in
arranged in 7 series. orders. 29 orders.
10. All flowering plants are placed in 280 families of flowering plants Number of recognised angio-
200 families. have been recognized. spermic families is 411.
11. Gramineae is the last family of Orchidaceae is the last family of Gramineae is the last family of
monocots. monocots. monocots.
primitive while those lacking subsidiary cells are advanced; (vi) Unilacunar nodes are derived from
trilacunar or pentalacunar nodes; (vii) Xylem fibres evolved from tracheids to libriform fibres, through
fibre tracheids; (viii) Cymose inflorescence is primitive while racemose is derived; (ix) Flowers with
an indefinite or a variable number of their floral parts are primitive; (x) Pollen grains with their exine
lacking any external sculpturing are primitive while those having various types of sculptures are
advanced; (xi) Apocarpous gynoecium is the characteristic of primitive taxa; (xii) Unitegmic ovules
developed from bitegmic ovules; (xiii) Basic type of ovule is anatropous type; all others are derived
ones; (xiv) Basic and most primitive type of female gametophyte is 8-nucleate Polygonum-type;
(xv) Primitive condition is porogamy, and the derived conditions are mesogamy and chalazogamy;
Classification 37
(xvi) Most primitive and basic type of fruit is a many-seeded follicle which develops from a multi-
carpellary apocarpous gynoecium.
Takhtajan divided Magnoliophyta (= Angiospermae) into two classes—Magnoliopsida (Dicots)
and Liliopsida (Monocots). He further divided Magnoliopsida into 7 subclasses, 20 superorders and
71 orders, and Liliopsida into 3 subclasses, 8 superorders, and 21 orders.
Takhtajan (1980) recognizes a total of 92 orders and 410 families among angiosperms. An outline
of his classification up to the level of orders is undermentioned:
DIVISION: MAGNOLIOPHYTA (ANGIOSPERMAE)
Class: Magnoliopsida (Dicotyledones)
Subclass 1. Magnoliidae
Superorder I. Magnolianae
Orders: 1. Magnoliales (Annonales), 2. Illiciales, 3. Laurales, 4. Piperales,
5. Aristolochiales.
Superorder II. Rafflesinae
Order: 6. Rafflesiales.
Superorder III. Nymphaeanae
Orders: 7. Nymphaeales, 8. Nelumbonales.
Subclass 2. Ranunculidae
Superorder IV. Ranunculanae
Orders: 9. Ranunculales, 10. Papaverales, 11. Sarraceniales.
Subclass 3. Hamamelididae
Superorder V. Hamamelidanae
Orders: 12. Trochodendrales, 13. Circidiphyllales, 14. Eupteleales, 15. Didymelales,
16. Hamamelidales, 17. Eucommiales, 18 Urticales, 19. Barbeyales, 20. Casuarinales,
21. Fagales, 22 Balanopales, 23. Leitneriales.
Superorder VI. Juglandanae
Orders: 24. Myricales, 25. Juglandales.
Subclass 4. Caryophylladae
Superorder VII. Caryophyllanae
Orders: 26. Caryophyllales, 27. Polygonales.
Superorder VIII. Plumbaginanae
Order: 28. Plumbaginales.
Subclass 5. Dilleniidae
Superorder IX. Dillenianae
Orders: 29. Dilleniales, 30. Paeoniales, 31. Theales, 32. Violales, 33. Begoniales,
34. Capparales, 35. Tamaricales, 36. Salicales.
Superorder X. Ericanae
Orders: 37. Ericales, 38. Ebenales, 39. Primulales.
38 Plant Taxonomy
Superorder XI. Malvanae

Orders: 40. Malvales, 41. Euphorbiales, 42. Thymeleales.
Subclass 6. Rosidae
Superorder XII. Rosanae
Orders: 43. Saxifragales, 44. Rosales, 45. Fabales, 46. Connariales, 47. Podostemales,
48. Nepenthales.
Superorder XIII. Myrtanae
Order: 49. Myrtales.
Superorder XIV. Rutanae
Orders: 50. Rutales, 51. Sapindales, 52. Geraniales, 53. Polygalales.
Superorder XV. Aralianae
Orders: 54. Cornales, 55. Apiales.
Superorder XVI. Celastranae
Orders: 56. Celastrales, 57. Santalales. 58. Balanophorales, 59. Rhamnales,
60. Elaeagnales.
Superorder XVII. Proteanae
Order: 61. Proteales.
Subclass 7. Asteridae
Superorder XVIII. Gentiananae
Orders: 62. Gentianales, 63. Oleales, 64. Dipsacales, 65. Loasales.
Superorder XIX. Lamianae
Orders: 66. Polemoniales, 67. Lamiales, 68. Scrophulariales.
Superorder XX. Asteranae
Orders: 69. Campanuales, 70. Calycerales, 71. Asterales.
Class: Liliopsida (Monocotyledones)
Subclass 1. Alismalidae
Superorder I. Alismatanae
Orders: 1. Alismatales, 2. Najadales.
Subclass 2. Liliidae
Superorder II. Triuridanae
Order: 3. Triuridales.
Superorder III. Lilianae
Orders: 4. Liliales, 5. Smilacales, 6. Burmanniales, 7. Orchidales, 8. Bromeliales.
Superorder IV. Juncanae
Orders: 9. Juncales, 10. Cyperales.
Superorder V. Commelinanae
Orders: 11. Commelinales, 12. Eriocaulales, 13. Restionales, 14. Hydatellales, 15. Poales.
Classification 39
Superorder VI. Zingiberanae

Order: 16. Zingiberales.
Subclass 3. Arecidae
Superorder VII. Arecanae
Orders: 17. Arecales, 18. Cyclanthales, 19. Pandanales, 20. Typhales.
Superorder VIII. Aranae
Order: 21. Arales.
Merits of the System of Takhtajan
1. Dicots (Magnoliopsida) are discussed prior to monocots (Liliopsida).
2. Dicots begin with the Magnoliales which are universally considered to be the most primitive
living angiosperms.
3. In this system the families are small homogeneous units made up of closely related
genera.
4. Division of Dicots into two traditional groups of Engler and Prantl, i.e. Archichlamydae and
Metachlamydae, has been abolished in this system.
5. Alismatales, which are considered to be the most primitive living monocots these days, are
the starting point of monocots in this system.
Demerit of the System of Takhtajan
One strong demerit is the extremely narrowly defined taxa in this system that has resulted in the
unwarranted splitting of the related groups.
3.6.2 Arthur Cronquist (1981)

On the basis of a wide range of taxonomic characters of phylogenetic importance, Cronquist (1981)
presented a phylogenetic classification. With the help of synoptic keys and detailed charts, he grouped
the angiospermic families into orders and subclasses on a worldwide basis. Pteridosperms (seed ferns)
were considered by him as the probable ancestors of angiosperms.
Class Magnoliopsida (= Dicots) in this system has been divided into six subclasses, i.e. Magnoliidae,
Hamamelidae, Caryophyllidae, Dilleniidae, Rosidae, and Asteridae. Of them, the Magnoliidae were
thought to be the basal complex, and all the remaining five subclasses were derived from it sepa-
rately. Asteridae formed the most advanced group of dicots.
Class Liliopsida (= Monocots) in this system has been divided into five subclasses, i.e. Alismatidae,
Arecidae, Commelinidae, Zingiberidae, and Liliidae. It is proposed by Cronquist (1981) that Liliopsida
have arisen from aquatic ancestors.
Cronquist (1981) recognises a total of 2 classes, 11 subclasses, 83 orders and 383 families and
about 2,19,300 species among the angiosperms. A brief outline of Cronquist’s system of classifica-
tion up to the level of orders is mentioned below. The number mentioned in the parenthesis of each
order signifies the number of the families in that order.
40 Plant Taxonomy
Division—MAGNOLIOPHYTA
Class—1. Magnoliopsida (Dicots)
Subclass 1. Magnoliidae
Orders: 1. Magnoliales (10), 2. Laurales (8), 3. Piperales (3), 4. Aristolochiales (1),
5. Illiciales (2), 6. Nymphaeales (5), 7. Ranunculales (8), 8. Papaverales (2).
Subclass 2. Hamamelidae
Orders: 1. Trochodendrales (2), 2. Hamamelidales (5), 3. Daphniphyllales (1), 4.
Didymelales (1), 5. Eucommiales (1), 6. Urticales (6), 7. Leitneriales (1), 8.
Juglandales (2), 9. Myricales (1), 10. Fagales (3), 11. Casuarinales (1).
Subclass 3. Caryophyllidae
Orders: 1. Caryophyllales (12), 2. Polygonales (1), 3. Plumbaginales (1).
Subclass 4. Dilleniidae
Orders: 1. Dilleniales (2), 2. Theales (18), 3. Malvales (5), 4. Lecythiadales (1), 5. Ne-
penthales (3), 6. Violales (24), 7. Salicales (1), 8. Capparales (5), 9. Batales
(2), 10. Ericales (8), 11. Diapensiales (1), 12. Ebenales (5), 13. Primulales
(3).
Subclass 5. Rosidae
Orders: 1. Rosales (24), 2. Fabales (3), 3. Proteales (2), 4. Podostemales (1),
5. Haloragales (2), 6. Myrtales (12), 7. Rhizophorales (1), 8. Cornales (4),
9. Santalales (10), 10. Rafflesiales (3), 11. Celastrales (11), 12. Euphorbiales
(4), 13. Rhamnales (3), 14. Linales (5), 15. Polygalales (7), 16. Sapindales
(15), 17. Geraniales (5), 18. Apiales (2).
Subclass 6. Asteridae
Orders: 1. Gentianales (6), 2. Solanales (8), 3. Lamiales (4), 4. Callitrichales (2),
5. Plantaginales (1), 6. Scrophulariales (12), 7. Campanulales (7), 8. Rubiales
(2), 9. Dipsacales (4), 10. Calycerales (1), 11. Asterales (1).
Class 2. Liliopsida (Monocots)
Subclass 1. Alismatidae
Orders: 1. Alismatales (3), 2. Hydrocharitales (1), 3. Najadales (10), 4. Triuridales
(2).
Subclass 2. Arecidae
Orders: 1. Arecales (1), 2. Cyclanthales (1), 3. Pandanales (1), 4. Arales (2).
Subclass 3. Commelinidae
Orders: 1. Commelinales (4), 2. Eriocaulales (1), 3. Restionales (4), 4. Juncales (2),
5. Cyperales (2), 6. Hydatellales (1), 7. Typhales (2).
Subclass 4. Zingiberidae
Orders: 1. Bromeliales (1), 2. Zingiberales (8).
Subclass 5. Liliidae
Orders: 1. Liliales (15), 2. Orchidales (4).
Classification 41
3.6.3 Rolf M.T. Dahlgren (1983)

Rolf Dahlgren (1980, 1981, 1983), Professor at the Botanical Museum of the University of Copenhagen,
Denmark, proposed a new classification on the basis of the characters drawn from several branches of
botany including anatomy, embryology, and chemistry. He believes that “none of the extant groups of
flowering plants is ancestral to any other present-day group. Thus, the Magnoliaceae-Ranunculaceae
assemblage is not ancestral to other flowering plants but has simply retained a large number of
primitive features” (Jones and Luchsinger, 1987). Dahlgren has been of the view that angiosperms
evolved in one particular line of gymnosperms, and are thus monophyletic. Magnoliiflorae have
been considered as most primitive dicotyledons while Alismatiflorae have been regarded as most
primitive monocots.
Dicots (Magnoliidae) have been divided into 25 superorders and 85 orders while monocots
(Liliidae) have been divided into 8 superorders and 25 orders. An outline of Dahlgren’s classification
up to the level of orders is mentioned below:
Class: MAGNOLIOPSIDA (= Angiospermae)
Subclass (1) MAGNOLIIDAE (Dicotyledoneae)
Superorder 1. Magnoliiflorae: Orders—l. Annonales, 2. Aristolochiales, 3. Rafflesiales,
4. Magnoliales, 5. Lactoridales, 6. Chloranthales, 7. Illiciales, 8. Laurales, 9. Nelumbonales.
Superorder 2. Nymphaeiflorae: Orders—l0. Piperales, 11. Nymphaeales.
Superorder 3. Ranunculiflorae: Orders—12. Ranunculales, 13. Papaverales.
Superorder 4. Caryophylliflorae: Order—14. Caryophyllales.
Superorder 5. Polygoniflorae: Order—15. Polygonales.
Superorder 6. Plumbaginiflorae: Order—16. Pumbaginales.
Superorder 7. Malviflorae: Orders—17. Malvales, 18. Urticales, 19. Euphorbiales,
20. Thymeleales, 21. Rhamnales, 22. Elaeagnales.
Superorder 8. Violiflorae: Orders—23. Violales, 24. Cucurbitales, 25. Salicales,
26. Tamaricales, 27. Capparales, 28. Salvadorales.
Superorder 9. Theiflorae: Orders—29. Dilleniales, 30. Paeoniales, 31. Theales.
Superorder 10. Primuliflorae: Orders—32. Primulales, 33. Ebenales.
Superorder 11. Rosiflorae: Orders—34. Trochodendrales, 35. Cercidiphyllales,
36. Hamamelidales, 37. Geissolomomatales, 38. Balanopales, 39. Fagales,
40. Juglandales, 41. Myricales, 42. Casuarinales, 43. Buxales, 44. Cunoniales,
45. Saxifragales, 46. Droserales, 47. Gunnerales, 48. Rosales.
Superorder 12. Podostemiflorae: Order—49. Podostemales.
Superorder 13. Proteiflorae: Order—50. Proteales.
Superorder 14. Myrtiflorae: Orders—51. Haloragales, 52. Rhizophorales, 53. Myrtales,
54. Chrysobalanales.
Superorder 15. Fabiflorae: Orders—55. Fabales.
Superorder 16. Rutiflorae: Orders—56. Sapindales, 57. Rutales, 58. Polygalales,
59. Geraniales, 60. Balsaminales, 61. Tropaeolales.
42 Plant Taxonomy
Superorder 17. Santaliflorae: Orders—62. Celastrales, 63. Vitales, 64. Santalales.

Superorder 18. Balanophoriflorae: Order—65. Balanophorales.
Superorder 19. Araliflorae: Orders—66. Pittosporales, 67. Araliales.
Superorder 20. Asteriflorae: Orders—68. Campanulales, 69. Asterales.
Superorder 21. Solaniflorae: Orders—70. Solanales. 71. Boraginales.
Superorder 22. Corniflorae: Orders—72. Fouquieriales, 73. Ericales, 74. Eucommiales,
75. Sarraceniales, 76. Cornales, 77. Dipsacales.
Superorder 23. Loasaliflorae: Order—78. Loasales.
Superorder 24. Gentianiflorae: Orders—79. Goodeniales, 80. Oleales, 81. Gentianales.
Superorder 25. Lamiiflorae: Orders—82. Scrophulariales, 83. Hippuridales, 84. Lamiales,
85. Hydrostachyales.
Subclass (2) LILIIDAE (Monocotyledoneae)
Superorder 1. Alismatiflorae: Orders—1. Hydrocharitales, 2. Alismatales 3. Zosterales.
Superorder 2. Triuridiflorae: Order—4. Triuridales.
Superorder 3. Ariflorae: Order—5. Arales.
Superorder 4. Liliiflorae: Orders—6. Dioscoreales, 7. Asparagales, 8. Liliales,
9. Burmanniales, 10. Orchidales.
Superorder 5. Bromelliflorae: Orders—11. Velloziales, 12. Bromeliales 13. Haemodorales,
14. Philydrales, 15. Pontederiales, 16. Typhales.
Superorder 6. Zingiberiflorae: Order—17. Zingiberales.
Superorder 7. Commeliniflorae: Orders—18. Commelinales, 19. Hydatellales, 20. Juncales,
21. Cyperales, 22. Poales.
Superorder 8. Areciflorae: Orders—23. Arecales, 24. Cyclanthales, 25. Pandanales.
3.6.4 Robert F. Thorne (1983)

On the basis of characters of comparative morphology, ultrastructure, embryology, palaeobotany,
cytology, plant geography, pollen and seed morphology, and host-parasite relationships, Thorne (1983)
published a pure phylogenetic system. He also believed in the monophyletic origin of angiosperms.
Thorne divided class Angiospermae (Annonopsida) into 2 subclasses, 28 superorders (with the
ending of iflorae), 54 orders, 73 suborders, 350 families, 12,255 genera and 2,25,490 species. A brief
outline of his classification up to the level of orders is mentioned below:
Class: ANGIOSPERMAE (Annonopsida)
Subclass 1. DICOTYLEDONEAE (Annonidae)
Superorder 1. Annoniflorae: Orders—1. Annonales, 2. Nelumbonales, 3. Paeoniales,
4. Berberidales.
Superoder 2. Nymphaeiflorae: Order—5. Nymphaeales.
Superorder 3. Rafflesiflorae: Order—6. Rafflesiales.
Superorder 4. Theiflorae: Orders—7. Theales, 8. Ericales, 9. Fouquieriales, 10. Ebenales,
11. Primulales, 12. Polygonales.
Classification 43
Superorder 5. Chenopodiiflorae: Orders—13. Chenopodiales, 14. Geraniales.

Superorder 6. Celastriflorae: Order—15. Celastrales.
Superorder 7. Santaliflorae: Orders—16. Santalales, 17. Balanophorales.
Superorder 8. Violiflorae: Orders—18. Violales, 19. Capparales.
Superorder 9. Malviflorae: Orders—20. Malvales, 21. Urticales, 22. Rhamnales,
23. Euphorbiales.
Superorder 10. Rutiflorae: Order—24. Rutales.
Superorder 11. Proteiflorae: Order—25. Proteales.
Superorder 12. Hamamelidiflorae: Orders—26. Hamamelidales, 27. Casuarinales, 28. Fagales.
Superorder 13. Rosiflorae: Orders—29. Rosales, 30. Pittosporales.
Superorder 14. Loasiflorae: Order—31. Loasales.
Superorder 15. Myrtiflorae: Order—32. Myrtales.
Superorder 16. Gentianiflorae: Orders—33. Oleales, 34. Gentianales, 35. Bignoniales,
36. Lamiales.
Superorder 17. Solaniflorae: Orders—37. Solanales, 38. Campanulales.
Superorder 18. Corniflorae: Order—39. Cornales, 40. Araliales, 41. Dipsacales.
Superorder 19. Asteriflorae: Order—42. Asterales.
Subclass 2. MONOCOTYLEDONEAE (Liliidae)
Superorder 1. Liliiflorae: Order—1. Liliales.
Superorder 2. Triuridiflorae: Order—2. Triuridales.
Superorder 3. Alismatiflorae: Orders—3. Alismatales, 4. Zosterales, 5. Najadales.
Superorder 4. Ariflorae: Order—6. Arales.
Superorder 5. Cyclanthiflorae: Order—7. Cyclanthales.
Superorder 6. Pandaniflorae: Order—8. Pandanales.
Superorder 7. Areciflorae: Order—9. Arecales.
Superorder 8. Typhiflorae: Order—10. Typhales.
Superorder 9. Commeliniflorae: Orders—l1. Commelinales, 12. Zingiberales.
Merits of Thorne’s System
1. Annonales are universally accepted as the most primitive living angiosperms, and they form
the starting point of this system.
2. Traditional division of dicots into Archichlamydeae and Metachlamydeae has been abolished
in this system.
3. In this system, by and large, the closely related taxa are placed nearer to one another.
4. The families of “Amentiferae” (a group of unrelated families) have been distributed in dif-
ferent orders, and the group Amentiferae has been abolished.
5. Inclusion of orders Malvales, Urticales, Rhamnales and Euphorbiales under one superorder
Malviflorae is also a commendable aspect of this system.
44 Plant Taxonomy
6. Placing of the related orders Cornales and Dipsacales in one superorder Corniflorae is also
the merit of this system.
Demerits of Thorne’s System
1. This system is not of much practical utility in the identification of plants.
2. The view of Thorne that angiosperms might have originated from some Pteridospermous
members in early Cretaceous times is also not accepted by several taxonomists.

1. What are the basic categories of systems of classification? Dioscorides, A.P. de Candolle,
and Bentham and Hooker proposed which type of system of classification?
2. Who is called the “father of taxonomy”?
3. The classical works entitled Systema Naturae, Genera Plantarum, and Species Plantarum,
have been authored by which taxonomist/taxonomists?
4. Linnaeus divided plants into 24 classes. Name at least 10 of the classes proposed by him.
5. Bentham and Hooker described ________ species belonging to ________ genera of
________ families of flowering plants in his Genera Plantarum.
6. Give an outline of the system of classification proposed by Bentham and Hooker.
7. Write some merits and demerits of the system of Bentham and Hooker.
8. Who has authored “Die Naturlichen Pflanzenfamilien”?
9. Hutchinson based his system of classification on how many principles? Write at least 10 of
these principles.
10. Give a comparison of the systems of Bentham and Hooker and Hutchinson.
Suggested Reading
Bentham, G. and J.D. Hooker, 1862–1883, Genera Plantarum, 3 Vols., London.
Bessey, C.E., 1915, Phylogenetic taxonomy of flowering plants, Ann. Mo. Bot. Gard. 2: 109–164.
Candolle, A.P. de., 1813, Theorie elementaire de Ia botanique, Paris.
________ 1824–1873, Prodromus systematics naturalis regni Vegetabilis, 17 Vols., Paris.
Cronquist, A., 1981, An Integrated System of Classification of Flowering Plants, Columbia University
Press, New York.
Dahlgren, R., 1980, A revised system of classification of angiosperms, Bot.J. Linn. Soc. 80: 91–124.
________ S. Rusendal-Jensen and B.J. Nielsen, 1981, A revised classification of the angiosperms with com-
ments on the correlation between chemical and other characters. In D.A. Young and D.S. Seigler (eds.),
Phytochemistry and Angiosperm Phylogeny, Praeger, New York.
________ 1983, General aspects of angiosperm evolution and microsystematics, Nord. J. Bot. 3: 119–149.
Classification 45
Eichler, A.W., 1875–1878, Bluthendiagramme construirt und erlautert, 2 Vols. Leipzig.

Engler, A. and K. Prantl, 1887–1915. Die Naturlichen Pflanzenfamilien, 23 Vols., Leipzig.
Hutchinson, J., 1964–1967, The Genera of Flowering Plants, 2 Vols., Clarendon, Oxford.
________ 1969, Evolution and Phylogeny of Flowering Plants, Academic Press, London.
________ 1973, The Families of Flowering Plants (3rd ed.), Clarendon, Oxford.
Jussieu, A.L. de., 1789, Genera Plantarum Secundum Ordines naturales disposita, Paris.
Linnaeus, C., 1735, Systema Naturae, Lugduni Batavorum.
________ 1737, Genera Plantarum, Lugduni Batavorum.
________ 1753, Species Plantarum, 2 Vols., Stockholm.
Takhtajan, A., 1980, Outline of the classification of flowering plants (Magnoliophyta), Bot. Rev., 46:
225–359.
Thome, R.F., 1976, A phylogenetic classification of the angiospermae, Evol. Biol., 9: 35–106.
________ 1983, Proposed new realignments in the angiosperms. Nord. J. Bot. 3: 85–117.
C
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TAXONOMIC
STRUCTURE 4
Subordination of organisms in groups under groups is a fact of nature, and it results from evolutionary
divergence. When an ancestral form splits into subgroups which undergo divergence during course
of time, the resultant descendants are modified forms but resemble each other in decreasing degrees.
Members of a species resemble each other because they are genetically more close to each other.
This resemblance, however, decreases progressively in different species of the same genus, different
genera of the same family, different families of the same order, different orders of the same class
and so on. And, to establish such relationships is one of the main concerns of plant taxonomy. In
any biological classification, all this is accomplished by taxonomic hierarchy. Different hierarchical
categories of plant taxonomy have been mentioned under Article 1.5 of Chapter 1 (Table 1.1).
4.1 CONCEPT OF TAXA

Taxon (pl. taxa) may be defined as “a named taxonomic group of any rank”. Thus at family level, taxa
may be represented by the Ranunculaceae and Rosaceae, while Ranunculus and Rosa are examples
of generic taxa. The term “taxon” was actually coined to replace clumsy phrases such as taxonomic
entity and taxonomic unit. Furthermore, the organisms contained within a rank (e.g., species, genus,
or order) can also be referred to as taxa.
4.1.1 Taxonomic Characteristics and Taxonomic Characters

Characters by which an organism or group of organisms can be recognised is called a characteris-
tic. For example, flowers are characteristic of angiosperms and wood is characteristic of trees. On
the other hand, any part or shape of an organism that makes it possible to classify the organism is
called a character. For example, characters used in classification include the shape of the leaves,
arrangement of the reproductive organs, etc.
Taxonomic characters are actually features, such as form, structure, behaviour, and physiology,
that are assessed in isolation from the rest of the plant by taxonomists, for making comparisons and
interpretations. It is important to differentiate between characters and character states. For example,
width of the leaf may be a character, while leaves 6 mm wide are an expression of that character,
i.e., its character state. Often, characters are referred to as “good” or “bad” but this is strictly relative.
Taxonomic Structure 47
Thus, a good diagnostic character, such as compound leaves in a group of plants that mainly possess
entire leaves, would be a bad character for separating taxa in a group in which leaf divisions were
either variable or often compound.
4.1.2 Problems of Hierarchy in Taxonomic Structures

It has been established that taxonomic structure is a hierarchical system. For the effective working
of this system, there should exist clear concepts regarding various taxa in plant taxonomy. Differing
from non-living things, a biologist faces peculiar type of problems in circumscribing the different
categories or taxa in biology. It is so because they show dynamic, evolving and variable popula-
tions. To solve this problem, any concept of taxa or taxonomic category has to be flexible to some
extent. It should be able to meet the special requirements of biology. Variability within and also in
between the groups has caused many subjective judgements. At higher levels of hierarchy, this has
also resulted in various types of controversies. For example, traditionally, there were two kingdoms,
viz., Animalia and Plantae. Such a classification has been challenged by several modern workers.
Copeland (1947) suggested four kingdoms, viz. Monera, Protista, Metaphyta and Metazoa. Whittaker
(1969) gave a five-kingdom classification of organisms, namely Monera, Protista, Plantae, Fungi and
Animalia. Edwards (1976) recognized seven kingdoms under two superkingdoms (Procaryota and
Eucaryota).
It has, however, been observed that the degree of arbitrariness in classification decreases as we
proceed down in the hierarchy, and it reaches to its minimum at a particular level i.e., the species.
4.2 CONCEPT OF SPECIES

A species is usually the smallest unit of classification. It includes individuals which are alike and
can breed with each other. Species are sometimes divided into subspecies and varieties on the basis
of small differences between populations. Species are given Latin binomial names.
Species is, thus, the fundamental unit of study in taxonomy, comprising all the populations of one
breeding group that normally are permanently separated from other such groups by marked disconti-
nuities. In case, the crossing between species does not occur then the resulting hybrids are normally
sterile, thus maintaining the reproductive barrier between species. This broadly genetic definition of
a species does not hold good for species that reproduce by self-fertilisation or by asexual means or
to extinct species. In all these cases, a species is delimited by observation of the similarities between
its members and dissimilarities between it and the other species.
With reference to botanical nomenclature, the specific epithet forms the second part of the binomial
(the first part being the generic name) and it is always written in lower case. In plants, the ending
of the specific epithet always agrees with the gender of the generic name.
There exist several infraspecific categories in the taxonomic hierarchy, but subspecies and then
the varieties are probably the most widely used. Groups of similar species are placed in genera.
4.2.1 Subspecies
A subspecies is a taxon within a species. All subspecies of a species differ in small ways. Although
they can breed with each other, they are usually found in different places or in different populations.
48 Plant Taxonomy
While naming a subspecies, a third Latin name is put after the binomial. Binomial is the Latin name
of a species consisting of two words, of which first one is the name of the genus to which the spe-
cies belongs and the second one is the name which distinguishes the species from other species in
the same genus.
Subspecies is, thus, the rank subordinate to species in the taxonomic hierarchy. The term ‘sub-
species’ is used when two or more populations are separated in some way (e.g. morphologically or
ecologically) throughout their range. However, they are not usually genetically isolated. Generally, if
90% or more of a group of infraspecific individuals are recognizably distinct from another similar
group, then each may be ranked as subspecies. Often, this is referred to as “90% rule”.
The abbreviation “ssp.” or “subsp.” is used to indicate a subspecies, e.g. Portulaca pilosa ssp.
pilosa and Daucus carota subsp. gummifer.
4.2.2 Variety
The variety is a rank subordinate to species but above the category “form” in the taxonomic hierarchy
(Table 1.1, Chapter 1). Varieties are actually morphological variants, which may or may not have
a clear geographical distribution. Sometimes they represent only habit phase or colour. A variety
designated by one author may be designated as a subspecies or form by another author. A variety
produced by agricultural or horticultural techniques and not normally found in natural populations
is called a cultivated variety or cultivar. In a variety, there may also be recognised subvarieties.
4.2.3 Form
“Form” is the lowest rank (see Table 1.1, Chapter 1) normally used by taxonomists for sporadic
distinct variants that sometimes occur in populations. Forms may be relatively minor genetic vari-
ants of a variety or subvariety but their effects can be conspicuous. Commonly observed forms are
those in which flower colour is modified, e.g., occurrence of albino individuals in a population of
purple-flowered plants. Although, a form is more commonly used to distinguish variants of subspe-
cies and varieties, it may also be related directly to a species. Sometimes, “subforms” may also be
recognised within a form.
Above-mentioned details show that a genus is a group of species and a subspecies or a variety
or a form are parts of a species. Workers like Levin (1969), Cock (1977) and Wiley (1980) have
defined “species” in their own ways but still the definition of the term ”species” is a matter of great
discussion and controversy. Various views have been put forward by the biologists during last few
centuries. For the sake of convenience these can all be classified into four major concepts, viz.,
(i) nominalistic concept, (ii) taxonomic concept, (iii) biological concept, and (iv) alternative concept
of species. All these are discussed below, in brief.
(a) Nominalistic Concept of Species
According to Slobodchikoff (1976), the nominalistic concept of species suggests that the “nature
produces the individuals and nothing more”. Species is nothing but the brainchild of man and are not
objectively real. Different species can be explained only in terms of formal relationship and not on
the characteristics of organisms. According to workers like Burma (1954) and Spurway (1955), species
have been invented as a device to refer to large number of individuals collectively. Evolutionists like
Haldane (1956) believe that species do not exist as taxa and that it is simply a unit of convenience.
About “species”, nominalists think that individuals are related and hence included in a species due to
similarity whereas the evolutionary point of view of workers like Mayr (1969, 1976) is that similarity
is due to evolutionary relationship, i.e., it is a common descent.
(b) Taxonomic or Typological Concept of Species
The name ‘species’ comes from the Latin root “specere” which means “to look at”. It, therefore, refers
to appearance. Aristotle, the great philosopher believed that biological species are highly variable and
also opined that hybridization between different species resulted into new species. Such examples
have also been given that different breeds of dogs have come into existence from breeding of bitches
with tigers, lions, wolves, foxes and even goats. Theophrastus, the Father of Modern Botany, wrote a
book “Enquiry into Plants” on transmutation of species. He opined that species changed when they
are transplanted in different soil and living in different climate. Several ancient workers reported that
orange was a new species created by grafting lemon on pomegranate. Similarly, banana originated
from the seed of date-palm (Zirkle, 1935) inserted into the corm of Colocasia.
(1) Aristotelian Concept of Essentialism Aristotle suggested that for every natural group, there is
an inner “essence” that makes them what they actually are and that this is real. This concept is
also known as the “typological concept of species”. According to this concept, as mentioned by
Simpson (1961), “every natural group of organisms, hence every natural taxon in classification has
an invariant, generalised or idealised pattern shared by all members of the group”. Species were
taken to be fixed units in nature, their number determinate and they were characterised by universal
types or “essence”. The essence is like floral diagram of a family, and all the genera are supposed
to fit in this and anything less or more is inconsequential. According to this concept, each species is
supposed to have a basic plan which is unchangeable and defined also by the universal characteristics
possessed by all members of the group. Several evolutionists refused to accept the taxonomic concept
of species because “they do not necessarily reflect actual species in nature.”
But, what exactly constitute the “essence” is a basic question. Supporters of the “concept of
essence” believe that for all practical purposes, the “essence” is deduced by observing a number of
individuals. The constant and invariable ones should constitute the “essence”.
(2) John Ray’s Concept of Species Botanists, such as John Ray (1686), put forward the theory of
“breeding relationship as the basis of species”. He also suggested that there exist variability within
species. In his Historia Plantarum published in 1686, Ray suggested “great care should be taken
in deciding what constitutes a species and what sort of characters are insufficient for species
delimitations”.
(3) View of Linnaeus Linnaeus (1707–1778) recognised two different kinds of variations among
organisms, the true differences created by the Creator and constituting the “essence” and the
intraspecific variations which were considered to be “sport of nature” or “accidents”. Linnaeus (1737)
warned that “sport” should not be neglected. “If neglected, these elusive ghosts glide away and are
gone”. Linnaeus, however, realised in the later years that species can arise by hybridisation.
(4) Darwin’s view Darwin (1859), however, provided a sound basis by explaining evolution of
species by natural selection. According to him, the organisms are characterised by variation. The
geometrical increase in the number of individuals takes place due to fertility of the organisms. But,
50 Plant Taxonomy
it is subject to nature check i.e. the natural selection. Amongst these individuals only those survive
which possess an inherent advantage over the others in the population. These inherent advantages
are inheritable. For thousands of generations, the selection continues and new variants take the place
of the original ones in a rapidly changing environment. Mechanism of heredity was, however, not
known much during the times of Darwin.
(5) Neo-Darwinism Rediscovery of Mendel’s work in 1900 brought forward a new theory of evolution
in the 20th century, after the death of Darwin. It is called neo-Darwinism. It includes Darwin’s
theory of natural selection and the more recent knowledge of genetics and inheritance through
chromosomes. Due to this theory, phylogenetic relationship came to be accepted as the basis of
organismal similarities. It has been suggested that groups of organisms are not related because they
are similar, but they are similar because they have a common descent.
(c) Biological Concept of Species
Concept of species has altogether changed during last century due to considerable amount of work
done on various aspects, such as (i) genetic basis of variations, (ii) reproductive mechanisms, (iii)
population structure, (iv) breeding behaviour, etc. All these have led to the formulation of biologi-
cal concept of species by Mayr (1969), which has been supported by workers like Stebbins (1970),
Heywood (1974), and others. Mayr (1969) defined the biological concept of species by genetic kin-
ship as evidenced by breeding behaviour. Grant (1959) defined such a species as “a community of
cross-fertilising individuals linked together by bonds of mating and isolated reproductively from
other species”. This shows that biological species is formed by “groups of interbreeding natural
populations that are reproductively isolated from such other groups”. Two points mainly considered
in biological concept of species are (i) interbreeding between the members of the same species, and
(ii) reproductive isolation between the members of different species. The biological concept of spe-
cies has received wide acceptance among the modern biologists and it has inspired several studies
on various aspects of biology, especially genetics and breeding behaviour.
One of the greatest shortcomings of biological species concept is that it is not applicable to the
non-sexual organisms.
(d) Alternative Concepts of Species
(i) Concept of Evolutionary Species Grant (1971) proposed the concept of evolutionary species which
is equally applicable both sexual and asexual organisms. This concept as defined by Grant represents
“a spatio-temporal lineage of populations that evolve separately from other lineages and has its
own ecological niche”. In its present form, however, this concept is too vague to be applied in all
forms.
(ii) Concept of Sibling Species In several plant and animal species, there exist many instances where
“morphologically similar or identical populations are reproductively isolated”. Such species have
been named as sibling species by Mayr (1963). According to Amadon and Short (1976), the term
“sibling species” is used as a “purely descriptive term having no genetic or taxonomic implication”.
Stayskal (1972) criticized the use of the term “sibling species” because it exactly means of “same
parentage”. Instead, he used the alternative term “aphanic species” for “sibling species”.
(iii) Concept of Ecological Species According to Van Velen (1976), “species are maintained for the
most part ecologically and not reproductively”, and this is called concept of ecological species. This
concept suggests that (i) genes are of minor importance in evolution, (ii) the control of evolution is
mainly by ecology and the constraints of individual development, and (iii) selection works primarily
on phenotypes which are the main building blocks of community. Van-Velen defined species as
“a lineage which occupies an adaptive zone minimally different from that of any other lineages
outside its range”.
(iv) Concept of Selection Species Slobodchikoff (1976) viewed species as a unit of selection and this
formed the basis of the concept of selection species. According to him, “species is a system of
genetically similar individuals and populations maintained as a cohesive unit by a set of selection
pressures that balance the disruptive forces imposed by environmental factor, mutations or genetic
recombinations”. The entire concept of Slobodchikoff turns round selection pressures.
Conclusion: All the above-mentioned concepts clearly indicate that there is no universally appli-
cable definition for species. In biology, the rank of species covers a variety of situations, and therefore,
the concept or definition of species should be excercised by considerable flexibility. To conclude, it
may be mentioned that species is a group of individuals that (1) actually or potentially interbreed
with each other but not with other such groups, (2) show continuous morphological variation within
the group but which is distinct from other such groups. Taxonomically, species are grouped into
genera and divided into subspecies and varieties or, horticulturally, into cultivars.
4.3 CONCEPT OF GENUS

Most widely accepted definition of the word “genus” (pl. genera) is that “it is a group of related spe-
cies”. In the taxonomic hierarchy, “genus” is an important rank which is subordinate to family, but
above the rank of species. It is a group of obviously homogeneous species. The generic name forms
the first part of the binomial (the second part being the specific epithet), e.g. Cicer arietinum.
Genus is usually a singular noun and is written in Latin with a capital initial letter. Collections
of similar genera are grouped into families. Davis and Heywood (1963) have suggested three main
parameters that should provide guidelines for the determination of the generic status, viz. (i) natu-
ralness, (ii) delimitation of closely related genera, and (iii) practicability of keeping them distinct or
including them in other genera.
The concept of monophyletic taxa has proved quite satisfactory in determining the naturalness
and demarkation between different genera. Monophyletic group means that its members must have
descended from a common ancestor and it must include all the descendants of that ancestor. Legendre
and Vaillancourt (1969) defined genus as a monophyletic group of species which occupies a definite
adaptive zone. According to them, all the species descending from a common ancestor have to be
included in the same genus. All non-monophyletic genera have to be abandoned because they do not
represent natural groups.
In the light of recent developments the circumscription and delimitation of the genera may also be
redefined. For example, the “Eugenia complex” of Myrtaceae may include all the new world species
in the genus Eugenia Linn. and all the old world species in the genus Syzygium Gaertn.
However, while revising the generic limits, it should be done mainly on the basis of a worldwide
study because it needs in-depth knowledge of the range and patterns of variations. Instead of taking
a single arbitrarily selected character for revising a genus, taxonomists should better be guided by
what is called a synthetic approach. More emphasis should be given on natural grouping.
52 Plant Taxonomy
4.4 CONCEPT OF FAMILY

A taxon consisting of related genera is called a family. The Latin names of families usually end
with “–aceae” e.g. Ranunculaceae, Malvaceae, etc. Eight exceptions to such an ending, however, are
Compositae, Cruciferae, Gramineae, Guttiferae, Labiatae, Leguminosae, Palmae and Umbelliferae.
International Code of Botanical Nomenclature has, however, proposed alternative names for these fam-
ilies as Asteraceae for Compositae, Brassicaceae for Cruciferae, Poaceae for Gramineae, Hypericaceae
for Guttiferae, Lamiaceae for Labiatae, Fabaceae for Leguminosae, Arecaceae for Palmae, and
Apiaceae for Umbelliferae. Many workers, however, still use older names.
A family is a major category in taxonomic hierarchy comprising groups of similar genera. Some
taxonomists believe that “families” represent the highest natural grouping. Groups of similar families
are placed in orders.
Larger families may be split into tribes. The tribe is, therefore, a rank subordinate to family but
superior to genus in the taxonomic hierarchy (see Table 1.1, Chapter 1). The term ‘tribe’ is applied
to assemblages of similar genera within large families. The Latin names of tribes have the ending
“-eae”, e.g. tribe Saniculeae in the family Umbelliferae or Apiaceae. Similar tribes may be grouped
together in subfamilies. Tribes may also be split into subtribes. The Latin names of subtribes have
the ending “-inae”.
Ideally, families have to be natural and monophyletic. The characters, by which families are
delimited vary with the groups, but they are of more obvious nature than that of genera and species.
Both vegetative and reproductive characters are used in delimiting families. For example, Cactaceae
is recognised by the cactoid habit of the members while insectivorous families (e.g. Nepenthaceae
and Droseraceae) are characterised by their distinct insect-trapping mechanisms. Compositae and
Umbelliferae are recognised by their characteristic inflorescences while Poaceae is recognised by
their fruits.
Two different types of families are definable and indefinable (Walters, 1961). The families which
are very homogenous and natural groups are called definable, e.g. Brassicaceae and Apiaceae. These
can be easily recognised from such other familier because they are homogenous natural groups
and their component genera are ill-defined. On the other hand, indefinable families include great
diversity of structure and are not as distinctive as the definable families. Most of the present day
plant families are indefinable. Their member genera differ among themselves so much that they can
be easily separated, as in Ranunculaceae. Genera of indefinable families are easily definable due to
clear morphological discontinuities.
Sometimes, the question of inclusion of a particular genus in a given family becomes largely a
matter of choice of taxonomist. For example, Nyctanthes is included by some taxonomists in Oleaceae,
by others in Verbenaceae and by still others in Nyctanthaceae.
4.5 TAXA ABOVE FAMILY LEVEL

Order, Class, Division, and Kingdom are the categories above the level of family used in taxonomic
hierarchy. They are also sometimes divided into their lower categories like suborder, subclass and
subdivision. One more category “superorder” has been included by Takhtajan (1969) to accommodate
one more evolutionary node. Ideally, all these groups should be monophyletic. Regarding evolution
of these higher taxa (Kubitzki, 1977), much information is, however, not available.
Order is a taxon consisting of families. The Latin names of orders usually end with “-ales”, e.g.
Rosales. However, some orders, which were erected prior to the compilation of the International Code
of Botanical Nomenclature, end in “-ae”, e.g. Tubiflorae, Glumiflorae. Groups of families, thought
to possess a degree of phylogenetic unity, are placed in an order.
Class is a taxon consisting of orders. In Botany, class is a taxonomic rank below division and
above order. The names of classes end in “-phyceae” in algae, -mycetes in fungi, or -opsida in other
plants.
Division is a major taxon, which is made up of classes. Three main divisions of land plants
are bryophytes, pteridophtes and spermatophytes. A division is the second highest category in the
taxonomic hierarchy, placed above the classes and below the kingdom. The Latin names of division
terminate in “-phyta”, e.g. Tracheophyta. In place of division, several botanists now use the term
phylum.
Kingdom, the largest of all the taxa, is actually the highest level in the hierarchy of taxonomic
ranks. In older systems, of classification, there are only two kingdoms, viz., plant kingdom and
animal kingdom. In some modern systems of classifications, fungi are considered in a separate
kingdom—Mycota. Similarly, some taxonomists prefer to place unicellular organisms in their own
kingdom, the Protista. Whittaker (1969) proposed a five-kingdom system of classification of living
organisms, namely, Monera, Protista, Plantae, Fungi and Animalia.

1. Give a brief account of concept of species.
2. Explain briefly the concept of taxa.
3. Give an account of problems of hierarchy in taxonomic structures.
4. Describe briefly the biological concept of species.
5. Differentiate between: (a) subspecies and variety, and (b) variety and form.
Suggested Reading
Amadon, D. and L.L. Short, 1976, Treatment of subspecies approaching species status. Syst. Zool. 25:
161–167.
Cock, A.G. 1977, Bernard’s symposium – the species concept. Bio. J. Linn. Soc. 9: 1–30.
Edwards, P. 1976, A classification of plants into higher taxa based on cytological and biochemical criteria.
Taxon 25: 529–54.
Grant, V. 1971, Plant speciation. New York.
Haldane, J.B.S. 1956, Can a species concept be justified? Syst. Assoc. Publ. 2: 95–96.
Heywood, V.H. 1974, Principles and concepts in the classification of higher taxa. Pl. Syst. Evol. Suppl.
1: 1–12.
54 Plant Taxonomy
Legendre, P. and P. Vaillancourt 1969, A mathematical model for the entities, species and genus, Taxon
18: 234–252.
Levin, D.A. 1969, The nature of plant species, Science 204: 381–384.
Mayr, E. 1969, The biological meaning of species. Biol. J. Linn. Soc. 1: 311–320.
________ 1976, Is the species a class or an individual? Syst. Zool. 25: 19.
Slobodchikoff, C.N. 1976. Concepts of species. Vol. III. Pennsylvania.
Van Valen L. 1976. Ecological species, multispecies and oaks. Taxon 25: 233–239.
Walters, S.M. 1961, The shaping of angiosperm taxonomy, New Phytol. 60: 74–84.
Wiley, E.O. 1980, Is the evolutionary species fiction? Syst. Zool. 29: 76.
C
H
A
P
T
PLANT E
R
COLLECTION
AND SPECIMEN
PREPARATION 5
Plant collection and field preparation of specimens are the fundamental aspects of study, training
and research in plant systematics. Herbarium1 specimens are the permanent records of plant species
of a particular place at a given time. Therefore, the plants should be carefully collected, selected,
and the herbarium specimens should be properly prepared and preserved.
5.1 WHICH TYPE OF SPECIMENS SHOULD BE COLLECTED?

The points to be kept in mind during a plant collection are:
1. Collect entire, vigorously growing typical specimens.
2. Select such individuals that represent almost all phases of the natural population.
3. Avoid collecting insect-damaged specimens.
4. Collect underground parts (e.g. roots, bulbs, rhizomes, tuber, etc.) of herbaceous
perennials.
5. Collect those specimens of flowering plants that contain flowers, fruits and seeds, because
keys are prepared mainly on the basis of these characters.
6. Specimens larger than the size of a single sheet, should be divided and pressed on a series
of sheets.
7. Collect plants with the leaves intact as different kinds of foliage prove helpful in
identification.
8. Collect the bark and wood samples of the woody plants.
9. Avoid collecting rare or uncommon plants. Never collect the only plant of a species at a
locality.
5.2 FIELD EQUIPMENT AND SUPPLIES

Some of the commonly used field equipment and supplies needed for plant collection and prepara-
tion of herbarium specimens are:
1
For details of some national and international herbaria and their upkeeping, refer Chapter 15.
56 Plant Taxonomy
1. Field Press It is made up of a pair of hardwood, metal or strong plywood frames of 12 by

18 inch.
2. Driers or Blotters These are the sheets of heavy blotting papers or of other moisture absorb-
ing material of 11 by 16 inch dimensions. Old newspapers are also used as driers.
3. Straps or Ropes Rope, sash cord, or a pair of strong web straps of about 4–5 feet length
are used to tighten the press.
4. Corrugate Ventilators Sheets of corrugated cardboard of 12 by 18 inch are used in between
the driers (= blotters) as ventilators when plants are dried by artificial heat. They provide
space for the passage of air through the press to remove moisture.
5. Field Notebook An indispensable item of a plant collector is a permanently bound, small
field notebook with ruled pages. It is used to record the full data (such as date, place, local-
ity, habitat, elevation, local name, collector number,1 etc.) of the collected plant. The data is
recorded in the field notebook in the field itself.
6. Digging and Clipping Tools Trowel, diggers, hammer, pruning shears, garden clippers, geo-
logical pick, and heavy sheath knife, etc. are all used for digging or clipping the plants.
7. String Tags These are made of waterproof material, and are used for labelling plants that
are not immediately pressed.
8. Vasculum Vasculum is a container made of tin or aluminium sheet and contains a hinged
light lid. Plants, which are not pressed in the field, are placed in a vasculum to preserve
their freshness for sometime.
9. Collecting Bags These are plastic bags, used as containers for fresh specimens.
10. Collection Bottles These are glass or plastic bottles with leakproof screw caps. These are
used for collecting small-sized material, to be preserved in liquid preservatives.
11. Liquid Preservative Formalin-aceticacid-alcohol (FAA)2 is the most common liquid pre-
servative used for anatomy materials. 6 : 3 : 1 mixture of chloroform, 95% ethyl alcohol and
glacial acetic acid is often used for cytology materials.
12. Hand Lens A 5 ¥ or 10 ¥ lens is used in the field for observation and identification.
13. Waxed Paper Sheets of waxed paper are used for pressing viscid or weak-looking plants.
14. Cardboard Storage Boxes These are the boxes or containers used to store dry materials.
15. Maps These are useful in the field for determining the localities for particular species.
16. Colour Charts These are used for determining the actual colour of the flower parts in the
field itself.
17. Camera and Film These are used for taking photographs of important plants in the field.
18. Other Field Equipment and Supplies Altimeter, compass, pocket knife, soft lead pencils,
insect repellant, portable plant drier, and seed envelopes are some other useful equipment
for plant collection.
1
A collector number is a numerical series starting with 1 and continuing throughout the lifetime of the collector.
2
Composition of FAA should be 70% ethyl alcohol (90 cc), formalin (5 cc) and glacial acetic acid (5 cc).
Plant Collection and Specimen Preparation 57
5.3 ORGANISATION OF THE FIELD PRESS

Following should be the organisation of the field press:
Hardwood or plywood press
Corrugate ventilator
Drier or blotter
10 sheets of torn newspapers (each sheet will contain one specimen)
Drier or blotter
Corrugate ventilator
Drier or blotter
10 sheets of torn newspapers (each sheet will contain one specimen)
Drier or blotter
Corrugate ventilator (continue the sequence until 100 specimens i.e. 10 groups of 10 newspapers,
are included)
Hardwood or plywood press.
5.4 HOW TO PRESS PLANT SPECIMENS?

Specimens, after being cut or dug, should be pressed as soon as possible. The specimen can be
placed carefully on a pressing sheet, i.e. a newsprint sheet or a blotter, taking care so that there is
no folding or overlapping of parts. The extra leaves or branches are removed, if necessary. Plants
too large to fit in the 11 ¥ 16 inch fold of a newspaper or a blotter may be bent into a V, N, or
M figure. The specimens should not protrude from the fold of the paper.
Specimens should be arranged in such a way so that some upper and some lower surfaces of
leaves are exposed. Flowers or inflorescences should be spread out thoroughly for a complete view.
A few flowers can be cut longitudinally and pressed, if possible. Each sheet should contain a col-
lector’s number which refers to the notes in the collector’s field notebook.
In the press, the specimens in the specimen paper are placed in between two driers or blotters.
A corrugate ventilator is often inserted before the next specimen papers and driers are added. The
usual sequence, as shown above under Section 5.3 (Organisation of the Field Press), is corrugate
ventilator, drier, sheets of specimens in newspaper, drier, corrugate ventilator, drier, and so on.
All the specimens are placed for pressing in such a way so that the entire bundle is ultimately of
almost uniform thickness in the middle and sideways. Once the plant specimens are arranged, the
press is tightly bound with ropes or straps to prevent wrinkling of the specimens. The press is now
ready for drying.
5.5 DRYING OF SPECIMENS

Specimens should be dried as rapidly as possible to get the best results. In the usual process of
drying, the press containing the specimens is placed in the sun. After about 24 hours, the press is
opened, and the specimens are placed in fresh blotters. Any last rearrangement of the plant parts
58 Plant Taxonomy
may be made at this time. The press, along with the specimens in fresh blotters, is now again bound
tightly. The wet blotters are dried in the sun for reuse. For three-four days the wet blotters or dri-
ers are changed daily, until the specimens are completely dried. The press becomes loose when the
plants are completely dried.
Artificial heat may also be used for the drying process. But the specimens are never to be dried
in an oven. In humid regions, or the rainy season, the plants may be dried in a drier. A drier is
made in the form of a wooden box, 3 feet in length and 18 inches in breadth, made up of 1/10 inch
thick boards. Five light bulbs of 60 watts are fitted inside for producing heat. Small openings are
made at the bottom of the box for the entry of air which gets heated and will dry the plants. In this
way, a press containing 100 specimens can be dried in 8–12 hours.
An excellent drying chamber may also be made by using a wooden or metal box with an open
top that will accommodate a press. The box is equipped with an electric heater with a fan.
5.6 MOUNTING OF SPECIMENS

The process, by which a dry specimen is attached to a mounting paper or a herbarium sheet, and
a label is affixed at the lower right corner of the sheet, is called mounting. The standard size of a
herbarium sheet is 28.75 by 41.25 cm. Heavy, hand-made cardsheets are used as herbarium sheets.
Before mounting, the dry specimens are dipped in a saturated solution of mercuric chloride in
ethyl alcohol to prevent infection of fungi, insects, etc. In place of mercuric chloride, lauryl pen-
tachlorophenate is also used, because the former is a virulent poison.
A good quality glue or paste is applied to the back of the specimens for affixing them onto the
herbarium sheets. Common animal glue is used in India for this purpose. Some of the good quality
glues, used in developed countries, include Wilhold 128, Elmer’s Glue-All, Swifts Z-5032, or Nicobon
B. Strips of gummed paper are used as an additional aid to hold heavy and woody specimens. Loose
parts (e.g. seeds, fruits, flowers, etc.) and dissected parts are placed in a paper packet and pasted on
the same herbarium sheet. Use of cellulose tapes or stiching of the plant parts with thread should
be avoided.
5.7 DEEP FREEZING METHODS

These days, deep-freezing methods are being adapted in some herbaria to prevent infection of fungi,
insects, etc. instead of mercuric chloride and other harmful chemicals.
5.7.1 Portable Freezer Unit

Recently, Davis and Gauthier (2008) of University of Idaho, Moscow, evaluated and found that a
portable refrigerator/freezer unit, powered by the 12 v DC connection in a vehicle, maintained a
constant and colder temperature than a similar-sized cooler filled with ice. They found it useful for
preserving the quality of seeds and cuttings of plants collected in the field.
Portable freezer units offer two main benefits over traditional ice coolers. First, “refrigeration
control offers temperature stability independent of exterior temperature without the worries about
ice-melting”. Second, such units “can function well in remote locations where replenishment of ice
may not be feasible”. These units help very effectively in collecting seeds which require the essential
Plant Collection and Specimen Preparation 59
maintenance of appropriate temperature. Engel MT 35 is a portable freezer developed by Engel USA,

Jupiter, Florida (USA). The main drawback of Engel MT 35 is “its need to remain inside the vehicle
when sampling, although longer cables could resolve this concern” (Davis and Gauthier, 2008).
5.7.2 Deep-freeze Arctic Vault

Recently, in the early months of 2008, thousands of seeds, plants and crop varieties from several
countries belonging to all continents, have been shipped and preserved in a vault in the Arctic Circle.
According to the organisers of the Arctic Circle, all these would be preserved here in deep-freezers
for thousands of years for future study and use. In this Norwegian-built Svalbard Global Seed Vault
(SGSV), more than 2 lakh crop varieties will be stored in the deep-freeze conditions in the com-
ing years. The seeds of several crops including rice, wheat, beans, sorghum, sweet potatoes, lentils,
etc. have already been shipped to the vault’s built quite deep under the Arctic permafrost in this
Norway’s Svalbard archipelago. A very large number of seeds and crop varieties have been given by
Consultative Group on International Agricultural Research (CGIAR), which maintains seeds of over 6
lakh plant varieties in its various centres located in different parts of the world. A major centre of this
organisation is in Mexico City which maintains a bank of over 1.5 lakh unique samples of wheat and
its relatives from more than 100 countries and also a very vast collection of maize. Seed duplicates
from CGIAR shipped to this deep-freeze Arctic Vault were provided by various agricultural research
centres located in Benin, Colombia, Ethiopia, India, Kenya, Mexico, Nigeria, Peru, Philippines and
Syria. These CGIAR collections are referred as “crown jewels” of international agriculture. They
include the world’s largest and most diverse collections of rice, wheat, maize and beans.
5.8 LABELLING OF SPECIMENS

A label, usually of 6.5 by 10.5 cm (Jones and Luchsinger, 1987) dimension, is pasted on the lower
right side of a herbarium sheet. It contains definite information about the specimen pasted on the
herbarium sheet. Ideally, it contains a miniature essay on the plant and its habitat. The paper of the
label should be of high rag quality. .
Following information should be incorporated in a label:
1. Heading, indicating the name of the institution or person, along with the name of the state
and country.
2. Family name.
3. Scientific name, i.e. name of genus, species along with authority.
4. Locality, indicating the place of collection with its latitude and longitude, and distance from
a well-known place.
5. Habitat, indicating vegetation type, moisture, soil, slope, etc.
6. Date of collection.
7. Field notes from the collector’s field notebook.
8. Collector, mentioning his/her name.
9. Collection number, which is a numerical series starting with 1 and continuing throughout
the lifetime of the collector.
60 Plant Taxonomy
10. Local or vernacular name.

11. Additional Information, such as colour of the flower and leaves, associated plants, bark,
economic uses, etc.
5.9 IDENTIFICATION OF SPECIMENS

Finally, pressed, dried and labelled specimens are identified with the help of appropriate floras,
manuals, and monographs. A duplicate specimen is to be retained before sending a specimen to the
specialist for identification.
For more details of identification of plants, refer to Chapter 7.

1. Make a list of at least 10 commonly used field equipment needed for plant collection and
preparation of herbarium.
2. What is the most commonly used liquid preservative used for plant materials?
3. What should be the organisation of the “field press”?
4. Write a note on pressing and drying of plant specimens.
5. What information should be incorporated in labelling of specimens?
6. Give an account of some deep-freezing methods.
Suggested Reading
Archer, W.A., 1950, New plastic aid in mounting herbarium sheets, Rhodora 52: 298–299.
Croat, T.B., 1978, Survey of herbarium problems, Taxon 27: 203–218.
Davis, A.S. and M.M. Gauthier, 2008, Portable refrigerator freezer provides stable temperature for plant
material collection, Native Plants Journal 9(1): 41–43.
Davis, P.H., 1961, Hints for hard pressed collectors, Watsonia 4: 283–289.
Lee, W.L., B.M. Bell and J.F. Sutton, 1982, Guidelines for Acquisition and Management of Biological
Specimens, Assoc. Syst. Collections, Lawrence, Kansas.
Savile, D.B.O., 1962, Collection and Care of Botanical Specimens, Canada Deptt. Agricul. Publication
No. 11, 13.
Smith, C.E., 1971, Preparing Herbarium Specimens of Vascular Plants, Agricultural Information Bulletin
No. 348, U.S. Govt. Printing Press, Washington, D.C.
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EXAMINATION
OF A
PLANT SPECIMEN 6
Since, taxonomy is essentially a practical activity, examination of a plant specimen is one of the
most important work of a taxonomist. Without this, nothing can proceed in plant taxonomy. This
may be done in field, garden and/or in the laboratory with the help of specimens of living plants
or of well-mounted plants.
6.1 EQUIPMENT NEEDED FOR EXAMINATION OF PLANT SPECIMEN

Very simple type of equipment is needed for examining plant specimen. Some of the commonly
required articles include a hand lens (10X and 15X), a sharp rajor, blades, a pair of dissecting needles,
a pair of forceps, collection bottles, polythene bags, dissecting microscope, compound microscope,
a good drawing pencil, eraser, notebook, some old newspapers, and a good plant identification
manual.
6.2 INSTRUCTIONS TO BE FOLLOWED WHILE EXAMINING A PLANT

SPECIMEN
While examining a plant specimen, certain guidelines should be followed for getting best possible
results. Some of such guidelines are mentioned below:
1. Plants should be examined in upwards succession, beginning with the roots and ending with
the flowers and fruits.
2. Examine the given plant first with the naked eye and then with the help of the hand-lens.
3. Study the flowers with great care, again in ascending succession, i.e., starting from flower-
stalk through the calyx, corolla and androecium upto the tips of stigmas.
4. Make a record of all possible characters before the flower is cut in any way.
5. Use the hand-lens for recording the details like the way the anthers open, the aestivation,
the placentation, etc.
6. At least two flowers should be sectioned with the help of rajor or sharp blade, one horizon-
tally and the other down the middle. The former is required to know the placentation and
62 Plant Taxonomy
also to complete the floral diagram. The latter is necessary to see if the ovary is superior
or inferior and also to enable a half-flower to be drawn.
7. In half-flower drawings, cut surfaces should be drawn first, indicated by double lines, and
the background filled in later.
8. Floral diagram and other drawings should be drawn quite large in size.
9. The parts of the drawings should be labelled as much as possible.
10. If the given specimen has flowers at different stages, all stages should be examined and
drawn as much as possible.
11. It is often easier to see the aestivation in horizontal sections of buds. But better placentation
can be seen in young fruiting stage.
12. Make the maximum use of available material in an intelligent way as much as possible.
13. Develop a habit of making keen and precise observation of the given plant specimen.
14. For practical taxonomic work, accurate recording of the information obtained and making
their neat and correct diagrams is also highly essential.
6.3 GUIDE TO THE EXAMINATION OF PLANT SPECIMEN

Undermentioned is the scheme which should be used for the examination, description and identifica-
tion of a given plant specimen.
6.3.1 Habit
(1) Is the plant a herb or woody?
If it is a herb, then:
is it annual, biennial or perennial?
is it erect, prostrate, trailing, twining or climbing?
is it succulent?
is it aquatic?
If it is woody, then:
is it a shrub or tree?
is it evergreen or deciduous?
(2) Any other peculiar habit such as epiphyte, saprophyte, parasite, etc.
6.3.2 Root
(1) Is it tap root or adventitious or fibrous root?
(2) Branched or unbranched.
(3) Whether it shows any other special form such as fusiform1, napiform, conical, tuberous,
nodulose, annulated or moniliform, etc.?
1
For explanation of hundreds of technical terms used in this chapter, refer to Chapter 19 “550 Terms of Plant
Description”.
Examination of a Plant Specimen 63
6.3.3 Stem
(1) Is it herbaceous or woody?
(2) Is it erect, prostrate, climbing or twining? In case of a climber, note the means of climbing
like tendril, hook, spine or any other outgrowth?
(3) Is it cylindrical or angular? If angular, then note the number of angles.
(4) Is it unbranched or branched? If branched, then note the mode of branching.
(5) Is it hollow or solid, jointed or unjointed?
(6) Is it smooth, waxy, hairy or spiny?
(7) Is it green or of any other colour?
(8) Also note any other special modifications such as:
(i) rhizome, tuber, bulb, corm.
(ii) Whether it is a runner, sucker, stolon or offset?
(iii) Is it a phylloclade or cladode?
(9) Also note features of bud and bark, if present.
6.3.4 Leaf
(1) Note arrangement, i.e., whether the leaf is alternate, opposite or whorled.
(2) Note attachment to stem, i.e., whether it is sessile or petiolate.
(3) Note stipules, i.e., whether stipulate or exstipulate.
(i) If stipulate, then note the type of stipules, i.e., scaly, free-lateral, adnate, interpetiolar,
ochreate, foliaceous, etc.
(4) Note whether leaf is simple or compound.
(i) If compound, then mention whether pinnately compound or palmately compound.
In case it is pinnately compound, then note whether it is unipinnate, bipinnate, tripinnate
or decompound.
In case the leaf is palmately compound, then note whether it is unifoliate, bi-, tri-, quadri-,
or multi-foliate.
(5) Note shape of leaf, i.e., whether it is acicular, linear, lanceolate, elliptical, oval, ovate, obo-
vate, oblong, rotund, cordate, reniform, oblique, spathulate, sagittate, hastate, cuneate, deltoid,
falcate, pedate or lyrate.
(6) Note margin of the leaf, i.e., whether it is entire, repand, sinuate, serrate, dentate, runcinate,
crenate, ciliate or spinous.
(7) Note apex of leaf, i.e., whether the apex is acute, acuminate, obtuse, cuspidate, truncate,
retuse, emarginate, mucronate or cirrhose.
(8) Note surface of leaf, i.e., whether it is rough, glabrous, glaucous, glutinose, spiny or hairy.
(9) Note venation of leaf, i.e., whether it is reticulate or parallel.
64 Plant Taxonomy
(i) If it is reticulate, then note whether it is unicostate or multicostate. If multicostate, then

also note whether it is convergent or divergent type.
(ii) If it is parallel, then note whether it is unicostate or multicostate. If it is multicostate,
then also note whether it is convergent or divergent type.
6.3.5 Inflorescence
(1) Note whether the inflorescence is racemose or cymose, or special types, or flowers are
solitary.
(2) If racemose, then note whether it is a raceme, spike, spikelet, panicle, catkin, spadix, corymb,
umbel or capitulum
(3) If cymose, then note whether it is a monochasial, dichasial or polychasial cyme.
(4) If special type, then note whether it is a cyathium, or verticillaster, or hypanthodium.
6.3.6 Flower
Note whether the flower is:
(i) bracteate or ebracteate, (ii) bracteolate or without bracteoles, (iii) sessile or pedicellate, (iv) com-
plete or incomplete, (v) unisexual or bisexual, (vi) dimerous, tri-, tetra- or pentamerous, (vii) hyp-
ogynous, perigynous or epigynous, and (viii) colour.
6.3.7 Perianth
A collective term used together for calyx and corolla when there is no distinction between the two, as
in monocotyledons. Note the number of tepals, i.e., parts of perianth, and whether they are free (poly-
phyllous) or fused together (gamophyllous). Also note the colour of tepals and their aestivation.
6.3.8 Calyx
(1) Of how many sepals, is it composed?
(2) Are the sepals free (polysepalous) or united above the base (gamosepalous)?
(3) Whether aestivation is valvate, twisted, imbricate, quincuncial or vexillary?
(4) Colour of sepals, generally green or of any other colour.
(5) Caducous or persistent.
6.3.9 Corolla
(1) Number of petals.
(2) Whether free (polypetalous) or united (gamopetalous).
(3) Aestivation, as noted in case of calyx.
(4) Shape of corolla, i.e., whether cruciform, caryophyllaceous, rosaceous, tubular, campanulate,
rotate, funnel-shaped, papilionaceous, bilabiate, ligulate or personate.
(5) Appendages, if any, present on corolla such as nectary, corona, spur, etc.
6.3.10 Androecium
(1) Number of stamens (write ‘indefinite’ if more than ten).
(2) Count the number of whorls of stamens and note whether or not some stamens are reduced
into staminodes.
(3) Note whether stamens are free or united.
(4) Note the nature of cohesion, i.e., whether monadelphous, diadelphous, polyadelphous, synge-
nesious or synandrous.
(5) Nature of adhesion, i.e., whether epipetalous, gynandrous or possess any other special
character.
(6) Whether stamens are alternipetalous or obdiplostemonous.
(7) Inserted or exerted.
(8) Also note whether filament is long, short or flattened. Whether they show didynamous or
tetradynamous condition?
(9) Monothecous or dithecous.
(10) Attachment of the anthers, i.e., whether basifixed, adnate, dorsifixed or versatile.
(11) Whether introrse or extrorse or laterorse?
(12) Also note the presence of appendages, if any, e.g., hair, scales, staminal corona, etc.
(13) Whether any disc is present outside or inside the stamens?
6.3.11 Gynoecium
Note the following:
(1) Number of carpels.
(2) Whether carpels are free (apocarpous) or fused (syncarpous).
(3) Superior, inferior or semi-inferior nature of ovary.
(4) Number of locules, i.e., unilocular, bilocular or multilocular.
(5) Number of ovules in each locule.
(6) Placentation type, whether marginal, axile, parietal, free-central, basal or superficial.
(7) Number of styles.
(8) Number, shape and any type of modification of stigma.
(9) Presence or absence of disc below the ovary.
Difference Between Gynoecium, Pistil and Carpel
The gynoecium is the female part of a flower consisting of one or more pistils. A pistil is the female
reproductive organ of a flower, consisting of ovary, style and stigma. Carpel is the female reproduc-
tive unit of a flower, consisting of the ovary with ovules. Most angiosperms have several carpels,
which are joined together at their margins to form the ovary.
66 Plant Taxonomy
How to Calculate the Number of Carpels in a Pistil?

For calculating the number of carpels in a pistil, proceed as follows:
(1) Observe Carefully the Outline of the Ovary
(i) If it is cleary asymmetrical in cross-section, the pistil is likely to be made up of only one
carpel.
(ii) If it is symmetrical, the pistil is likely to be made up of two or more carpels. In case it is
symmetrically lobed into two or more lobes, the number of lobes may indicate the number
of carpels.
(2) Now Observe Carefully the Styles
(i) If the number of styles is more than one, then the pistil is made up of two or more carpels,
indicating that number of styles indicate the number of carpels.
(ii) In case there is only one style, the pistil may be made up of one or more than one carpels.
In such a condition, proceed as mentioned below in (3).
(3) Observe Carefully the Stigmas
(i) If there are present more than one stigmas, the pistil is made up of two or more carpels. If
there is only one stigma, then pistil may be made up of one or more than one carpel.
(ii) If the stigma is asymmetrical or horse-shoe shaped, it is made up of only one carpel.
(iii) If the stigma is symmetrical, the pistil is more likely to be made up of two or more
carpels.
(iv) If the stigma is lobed symmetrically into two or more lobes, the pistil is more likely to be
made up of two or more carpels. This indicates that the number of stigma lobes may indicate
the number of carpels.
(4) If the Stigma is Unlobed, then Proceed as Under
Cut a transverse section of the ovary across the middle and note whether the ovary is divided into
two or more compartments or loculi? If so, then the pistil is made up of two or more carpels,
indicating that the number of loculi indicate the number of carpels. If not, then pistil may be made
up of one or even more than one carpels.
(5) Observe the Number of Placentae Visible Clearly in a Cross-section
(a) If placentae are more than one, then the pistil is made up of two or more carpels, indicating
that the number of placentae indicates the number of carpels.
(b) If only one placentum is present, then pistil is more likely to be made up of only one
carpel.
By using a combination of the above-mentioned five methods, it is possible to ascertain if the
pistil is simple or compound.
(i) If the pistil is simple, then gynoecium is apocarpous and made up of a single free carpel.
(ii) If the pistil is compound, then gynoecium is syncarpous and made up of two or more fused
carpels.
In the syncarpous condition, also note the numbers of styles, stigmas and loculi in the ovary,
which together decide about the number of carpels making up the pistil.
6.3.12 Fruit
Record the type of fruit, if present, i.e., whether it is simple, aggregate or multiple type.
(1) If simple, then note whether it is dry or fleshy.
(a) If dry and simple, then note whether it is indehiscent (e.g., achene, utricle, caryopsis,
cypsella, or nut) or dehiscent (e.g., follicle, legume, siliqua, silicula or capsule) or schizo-
carpic (e.g., cremocarp, cacervulus, regma, lomentum or samara).
(b) If fleshy and simple, then note whether it is berry, hesperidium, pepo, drupe or pome.
(2) If aggregate, then note whether it is etaerio of achenes, or etaerio of follicles, etaerio of
drupes or etaerio of berries.
(3) If it is a multiple fruit, then note whether it is a sorosis or syconus.
6.3.13 Seed
Record the following:
(1) Number of seeds in a fruit.
(2) Whether it is endospermic or non-endospermic.
(3) Number of cotyledons.
(4) Shape, size and surface ornamentations of the seed.
6.3.14 Floral Formula

Write the floral formula of the given specimen according to the details given in Chapter 17 and
Table 17.1.
6.3.15 Floral Diagram

Make a floral diagram of the given specimen according to the guidelines discussed in Chapter 17.

1. What major guidelines one should follow while examining a plant specimen?
2. What should you note in a leaf while examining a plant specimen?
3. Make a systematic list of characters you should note in the male parts of a flower while
examining a plant.
4. How can you calculate the number of carpels in the pistil of a flower?
5. From the point of view of examining a plant, how should you study its fruit?
68 Plant Taxonomy
Suggested Reading
Featherly, H.I. 1954, Taxonomic Terminology of the Higher Plants, Iowa State College Press, Ames.
Lawrence, G.H.M. 1951, Taxonomy of Vascular Plants, Macmillan Company, New York.
Sharma, O.P. 2007, A Manual of Practical Botany Vol. II (9th ed.), Pragati Prakashan, Meerut.
Stearn, W.T. 1983, Botanical Latin (3rd ed.), Davis and Charles, Newton Abbot, England.
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IDENTIFICATION 7
7.1 WHAT IS IDENTIFICATION?
Identification is the basic process of classification. But it is not the same as naming of an individual.
In the biological sense, identification is the determination of the group to which a specimen belongs.
The process of identification usually includes a direct comparison of an unknown specimen with
the already classified, circumscribed, and named taxa. This process also includes the use of keys of
various types, computer-based methods and variety of polyclave-type devices.
In brief, identification may variously be defined as (i) the determination of similarities or dif-
ferences between two specimens; (ii) the direct comparison of the features of a specimen in hand
with those in keys in order to arrive at a name; (iii) the assignment of an unidentified taxon to
the correct class in an established system of classification; (iv) the determination of a name for a
particular specimen in relation to an already established system of identification.
7.2 CHARACTERS OFTEN CONSIDERED BEFORE PLANT IDENTIFICATION

1. Determine herbaceous or woody, and annual or perennial nature of the specimen.
2. Note the leaf type, phyllotaxy, and venation.
3. Note the presence or absence and type of stipule on young shoots.
4. Whether or not milky or coloured sap is present in the leaf, stem or other plant parts.
5. Note the distribution and kinds of surface coverings (i.e. hairs, trichomes, spines, etc.).
6. Observe the flower and name its parts.
7. Count the number of sepals and petals, and also note their arrangement, i.e. aestivation.
8. Note whether perianth is present in one series, more series, or absent.
9. Observe whether the sepals and petals are separate or fused.
10. Note whether pappus (e.g. Asteraceae) or epicalyx (e.g. Malvaceae) or similar structures are
present.
11. Note whether a nectar-secreting disc is present in the flowers (e.g. Rutaceae).
70 Plant Taxonomy
12. Determine whether the flowers are actinomorphic or zygomorphic.

13. Observe the number and attachment of stamens. Also note if there is any fusion of anthers
or filaments.
14. Note whether the stamens are antipetalous (e.g. Chenopodiaceae and Amaranthaceae), alter-
nipetalous or obdiplostemonous (e.g. Caryophyllaceae).
15. Count the number of pistils, styles and stigmas of the gynoecium.
16. Cut a transverse section of the ovary, count the number of locules, number of ovules per
locule, and also observe the placentation.
17. Cut a longitudinal section of the entire flower through its centre. Note the position of the
ovary and fusion of the perianth.
18. Determine the number of carpels.
After carefully examining all the above characters, the next step in identification is keying.
7.3 IDENTIFICATION WITH KEYS

A key is an artificial arrangement or analytical device whereby a choice is provided between two
contradictory statements resulting in the acceptance of one and the rejection of the other. A single
pair of contradictory statements in a key is called a couplet. Each statement of a couplet is termed
a lead. Leads are usually the best contrasting characters. Characters following the lead are called
secondary key characters.
Keys are of two types: punched cards keys and dichotomous keys.
7.3.1 Punched Cards Keys

These consist of cards having names of all the taxa (i.e. all species, genera or families for which
the key is meant) printed on all of them. On one of the corners of each card is printed any one
character and a definite number. All the taxa showing this particular character are indicated by a
perforation in front of their names, and the taxa lacking this character are without any perforation.
The number of the cards and the characters chosen for the purpose are the same. For identifying
a plant, only those cards showing characters possessed by this plant, are selected. Combination of
characters shown by this plant will allow only one perforation in the selected set of cards. The plant
is then referred to that particular family to which the card shows this perforation.
7.3.2 Dichotomous Keys

These keys consist of pairs of contrasting characters or couplets, each statement of which is a lead.
Both the leads are numbered, and begin with the same word as far as possible.
Dichotomous keys are of two general types i.e. indented key and bracket key.
(a) In the indented or yoked key, each of the couplets is indented a fixed distance from the left
margin of the page. An example of the indented key is given below in the form of identifi-
cation of 5 common genera of Ranunculaceae (Buttercup family), i.e, Ranunculus, Clematis,
Anemone, Aquilegia and Delphinium:
Plant Identification 71
Fruit a group of achenes; unspurred flowers.

Petals absent
Sepals usually 4; involucre absent .........................................................................Clematis
Sepals usually 5; involucre present....................................................................... Anemone
Petals present ......................................................................................................... Ranunculus
Fruit a group of follicles; spurred flowers.
Spurs 5; flowers regular ........................................................................................Aquilegia
Spur 1; flowers irregular ................................................................................... Delphinium
(b) In the bracket or parallel key, the two couplets are always next to each other in consecutive
lines on the page. At the end of each line in the key, there is either a number or a name refer-
ring to a couplet. An example of the bracket key is given below, in which all the same five
genera of Ranunculaceae (i.e. Ranunculus, Clematis, Anemone, Aquilegia and Delphinium)
are identified:
(1) Fruit a group of achenes; unspurred flowers ................................................................ (2)
(1) Fruit a group of follicles; spurred flowers .....................................................................(4)
(2) Petals absent ..........................................................................................................(3)
(2) Petals present.......................................................................................... Ranunculus
(3) Sepals usually 4; involucre absent ........................................................................Clematis
(3) Sepals usually 5; involucre present ..................................................................... Anemone
(4) Spurs 5; flowers regular ............................................................................Aquilegia
(4) Spur 1; flowers irregular ....................................................................... Delphinium
7.4 SOME UNCONVENTIONAL IDENTIFICATION METHODS

Keys are the conventional or traditional methods of identification. Some highly improved, recent but
unconventional methods of identification include polyclave identification, computerised identification
and other identification methods.
7.4.1 Polyclave Identification

Polyclave method is a system of identification in which a choice of several characteristics can be
used. A polyclave is actually a multientry, order-free key implemented in several different formats.
In a polyclave method of identification the user of the system is free to choose any character, in any
order or sequence, and thus avoid the rigid format of traditional dichotomous keys.
One form of the polyclave is a diagnostic key in which cards are utilised. The cards are placed
on top of one another to eliminate taxa which disagree with the plant to be identified. The second
form of the polyclave is a computer-stored multientry key, and the third form is a printed table which
gives the status of different taxa and characters useful for separating the taxa.
7.4.2 Computerised Identification

Computers are machines planned only for the evaluation of a computer programme or algorithm.
An algorithm is a series of logical steps or instructions by which an identification can be made.
72 Plant Taxonomy
Therefore, the computer can be programmed to identify only if an initial algorithm has been prepared
successfully by an expert systematist.
In the computerised identification of plants, the major research efforts are grouped into four major
approaches: (1) computer-constructed keys, (2) computer-stored dichotomous keys, (3) automated
pattern-recognition systems, and (4) simultaneous character-set methods.
A system of computer programme to help in the plant identification and other related aspects of
plant taxonomy has been presented by Morse (1974). This programme of Morse includes routines for
plant identification, key construction, description preparation, taxa comparison and also the produc-
tion of punched-card field keys.
7.4.3 Other Identification Methods

Some other unconventional, but sometimes quite effective, methods of plant identification include the
use of techniques of chromatography, spectroscopy and optical scanners, etc. Statistical methods
are also in the current use, in some developed countries, for plant identification.
7.5 ARTIFICIAL KEYS FOR THE IDENTIFICATION OF 51 COMMON FAMILIES

OF ANGIOSPERMS
7.5.1 KEY-1 (Major Groups)
1. Plants with seeds; ovules enclosed within the ovary ............................................ Angiosperms
2. Flowers mostly tetra- to pentamerous; two cotyledons ..................................Dicotyledons
3. Both sepals and petals present.
4. Petals are free; flowers with calyx and corolla .......................................... Polypetalae
5. Polysepalous; petals hypogynous.
6. Dome-shaped, unexpanded thalamus ............................................Thalamiflorae
6. Thalamus expanded into a disc..........................................................Disciflorae
5. Gamosepalous; inferior ovary ................................................................Calyciflorae
4. Petals are fused; flowers with calyx and corolla ..................................... Gamopetalae
7. Ovary inferior ..................................................................................................Inferae
7. Ovary superior
8. More than two carpels ..................................................................... Heteromerae
8. Only two carpels ............................................................................. Bicarpellatae
3. Either sepals or petals are present; rarely both absent ................................... Monochlamydae
2. Flowers mostly trimerous; one cotyledon.................................................. Monocotyledons
1. Plants with seeds; ovules exposed, i.e. not enclosed within the ovary .............. Gymnosperms
7.5.2 KEY-2 (Thalamiflorae)

DICOTYLEDONS
Polypetalae
Thalamiflorae
Dome-shaped unexpanded thalamus; polysepalous; hypogynous; ovary superior.
1. Carpels free (apocarpous).
2. Herbs or soft-wooded climbing shrubs ........................................................ Ranunculaceae
2. Woody shrubs or trees; rarely climbing
3. Leaves stipulate ......................................................................................... Magnoliaceae
3. Leaves exstipulate......................................................................................... Annonaceae
1. Carpels fused (syncarpous)
4. Parietal placentation.
5. Flowers tetramerous.
6. Androgynophore present .............................................................. Capparidaceae
6. Androgynophore absent; anthers tetradynamous ................................Cruciferae
5. Flowers pentamerous.
7. Stamens indefinite; flowers actinomorphic .................................. Papaveraceae
7. Stamens 5; flowers zygomorphic ......................................................... Violaceae
4. Axile placentation; calyx valvate; stamens monadelphous ...........................Malvaceae
4. Axile placentation; calyx valvate; stamens in different whorls ..................... Tiliaceae
4. Free-central placentation; stamens 10; obdiplostemonous ..................Caryophyllaceae
7.5.3 KEY-3 (Disciflorae)

DICOTYLEDONS
Polypetalae
Disciflorae
Thalamus expanded into a disc; ovary superior.
1. Leaves alternate; pinnately compound, gland-dotted; stamens not monadelphous... Rutaceae
1. Leaves are not gland-dotted; stamens monadelphous .............................................. Meliaceae
7.5.4 KEY-4 (Calyciflorae)

DICOTYLEDONS
Polypetalae
Calyciflorae
Calyx gamosepalous; thalamus cup-shaped; disc thin, often lining the calyx tube.
1. Calyx inferior; generally hypogynous to perigynous; leaves stipulate.
2. Leaves pinnately compound; calyx with odd sepal anterior; fruit a pod .....Leguminosae
74 Plant Taxonomy
3. Corolla descending-imbricate; flowers zygomorphic.........Fabaceae or Papilionaceae

3. Corolla ascending-imbricate; flowers zygomorphic............................. Caesalpinoideae
3. Corolla valvate; flowers actinomorphic .................................................... Mimosoideae
2. Calyx with odd sepal posterior; fruit of various types but not a pod ................Rosaceae
1. Calyx superior; flowers epigynous; leaves exstipulate.
4. Shrubs or trees.
5. Leaves gland-dotted and exstipulate ..............................................................Myrtaceae
4. Herbs.
6. Mostly climbers; flowers unisexual ........................................................ Cucurbitaceae
7. Mostly herbs and not climbers; flowers bisexual ..................................... Umbelliferae
7.5.5 KEY-5 (Inferae)

DICOTYLEDONS
Gamopetalae
Inferae
Petals fused; ovary inferior.
1. Leaves opposite or whorled; stamens epipetalous; dichasial cyme inflorescence ...................
Rubiaceae
1. Leaves alternate or opposite; stamens syngenesious; head or capitulum inflorescence ..........
Asteraceae or Compositae
7.5.6 KEY-6 (Bicarpellatae)

DICOTYLEDONS
Gamopetalae
Bicarpellatae
Petals fused; flowers with calyx and corolla; ovary superior; two carpels.
1. Flowers actinomorphic.
2. Leaves opposite; milky sap present.
3. Stamens 5, epipetalous; axile placentation; gynostegium absent .............Apocynaceae
3. Stamens 5, connected with stigma to form gynostegium; marginal placentation ........
Asclepiadaceae
2. Leaves alternate; milky sap generally absent.
4. Plants erect.
5. Carpels obliquely placed; swollen placenta ........................................... Solanaceae
5. Carpels not obliquely placed; placenta not swollen ...........................Boraginaceae
4. Plants climbing or prostrate; carpels medianly placed ........................ Convolvulaceae
1. Flowers zygomorphic.
6. Bracts large, leafy; stem with swollen joints; seeds with jaculators ............. Acanthaceae
6. Bracts minute or absent; stem not with swollen joints; seeds not with jaculators.
7. One or two ovules in each locule.
8. Ovary entire, style terminal, stigma simple......................................... Verbenaceae
8. Ovary entire, style long, stigma bifid ................................................... Pedaliaceae
8. Ovary tetralocular, style gynobasic .............................................................Labiatae
7. Many ovules in each locule................................................................ Scrophulariaceae
7.5.7 KEY-7 (Monochlamydae)

DICOTYLEDONS
Monochlamydae
Flowers incomplete; either calyx or corolla or sometimes both absent.
1. Flowers bisexual.
2. Leaves stipulate, stipules ochreate ..................................................................Polygonaceae
2. Leaves exstipulate.
3. Leaves opposite, bracts coloured ............................................................. Nyctaginaceae
3. Leaves alternate.
4. Flowers with dry, scarious bracts and perianth .............................. Amaranthaceae
4. Perianth sepalloid or green ............................................................. Chenopodiaceae
1. Flowers unisexual.
5. Inflorescence cymose or raceme; ovary unilocular ........................................... Urticaceae
5. Inflorescence cyathium; ovary trilocular ..................................................... Euphorbiaceae
5. Inflorescence catkin; fruit capsule; 2–4 stigmas ................................................Salicaceae
5. Inflorescence catkin; fruit drupe or sorosis; 2 stigmas.......................................Moraceae
7.5.8 KEY-8 (Monocotyledons)

Monocotyledons
1. Arborescent; stem with prominent scars of leaf bases; flowers in fleshy spikes ...................
Arecaceae or Palmaceae
1. Not arborescent; stems with no prominent leaf bases.
2. Herbaceous, aquatic; apocarpous ........................................................................Alismaceae
2. Plants generally terrestrial; usually syncarpous.
3. Inferior ovary.
4. Inflorescence spadix ...................................................................................Musaceae
4. Inflorescence not spadix.
5. Parietal placentation ......................................................................... Orchidaceae
5. Placentation usually axile, not parietal.
6. Six fertile stamens.................................................................. Amaryllidaceae
76 Plant Taxonomy
6. Only one fertile stamen; other stamens transform into staminodes .............
Zingiberaceae
3. Superior ovary.
7. Well-developed perianth.
8. Leaves with sheathing base ........................................................ Commelinaceae
8. Leaf bases not sheathing ....................................................................... Liliaceae
7. Perianth ill-developed or reduced to hairs or bristles.
9. Flowers arranged on spadix, fruit berry .................................................Araceae
9. Flowers arranged in spikelets; fruit indehiscent.
10. Phyllotaxy 1/3; culm triangular; perianth reduced to bristles or hairs;
fruit nut ....................................................................................... Cyperaceae
10. Phyllotaxy 1/2; culm cylindrical; perianth reduced to lodicules; fruit
caryopsis.....................................................................Poaceae or Graminae

1. Differentiate between punched card keys and dichotomous keys.
2. Give an account of dichotomous keys and their types with reference to plant identification.
3. Make an artificial key to differentiate major groups of dicotyledons.
4. How can you differentiate between Rubiaceae and Asteraceae making an artificial key?
5. Using an artificial key, differentiate between any four families of your course belonging to
Thalamiflorae.
Suggested Reading
Bossert, W., 1962, “Computer techniques in systematics”, In Systematic Biology, National Academy of
Science Publication, Washington.
Hansen, B. and K. Rahn, 1969, Determination of angiosperm families by means of a punched-card system,
Dan. Bot. Ark. 26: 1–46.
Harrington, H.D. and L.W. Durrel, 1957, How to Identify Plants? The Shallow Press, Chicago.
Hutchinson, J., 1967, Key to the Families of Flowering Plants of the World, Clarendon Press, Oxford.
Morse, L.E., 1971, Specimen identification and key construction with the time-sharing computer, Taxon
20: 269–282.
________ 1974, Computer-assisted storage and retrieval of the data of taxonomy and systematics, Taxon
23: 29–43.
Osborne, D.V., 1963, Some aspects of the theory of dichotomous keys, New Phytol. 62: 144–160.
Pankhurst, R.S.. 1970, A computer programme for generating diagnostic keys, Computer J. 13: 145–151.
________ 1971, Botanical keys generated by computer, Watsonia 8: 357–368.

________ 1974, Automated identification in systematics, Taxon 23: 45–51.
________ 1975, Biological identification with Computers, Academic Press, London.
Saldanha, C.J. and C.K. Rao, 1975, A punched-card key to the Dicot Families of South India, Amarind
Publishers, Bangalore.
C
H
A
PLANT P
T
NOMENCLATURE, E
R
BOTANICAL
NAMES AND
PHYLOCODE 8
A PLANT NOMENCLATURE
8.1 FUNDAMENTALS OF PLANT NOMENCLATURE

8.1.1 Definitions
Assignment of definite names to plants is called plant nomenclature.
In the present botanical world, the nomenclature involves the principles governed by rules for-
mulated and adopted by International Botanical Congresses. The rules developed by IBC are listed
formally in a code called International Code of Botanical Nomenclature (Voss et al., 1983), abbrevi-
ated as ICBN. The major goal of ICBN is to provide one correct name for each taxon.
Taxa (singular, taxon) are the taxonomic groups of any rank. The ascending hierarchy of taxa
include species, genus, family, order, class and division.
8.1.2 Who is a Nomenclaturist?

A nomenclaturist is a taxonomist who assigns names to new taxa, determines the right names for
old taxa according to the rules of International Code of Botanical Nomenclature, and finalises the
right name for a specimen according to an established system of classification.
8.1.3 Why is Nomenclature Needed?

Imagine for a few moments about the state of affairs if there are no names for all the things we see,
make or handle. How strange and chaotic the life would be in such a condition? The entire business
affairs of the world would stop, and practically there will be no “give and take” of knowledge. It
would also be impossible for all of us to communicate our feelings to others. Practically, the life
would stop. Therefore, nomenclature for everything is needed.
Plant Nomenclature, Botanical Names and Phylocode 79
8.2 COMMON NAMES AND SCIENTIFIC NAMES

Vernacular or common names are made up of words from the native language of the country or the
region. They may vary in different countries as well as in different regions of the same country. They
may be Portuguese words in Portugal; Spanish in Spain; English in England, U.S.A. and Australia;
and Hindi, Bangla, Oriya or Tamil, etc. in U.P., West Bengal, Orissa and Tamilnadu, respectively.
‘Papita’, ‘Kela’, ‘Gazar’ and ‘Tamatar’, of India are known in U.S.A. as papaya, banana, carrot and
tomato, and by other names in Spain. There is therefore no international uniformity in common
names.
On the other hand, scientific names which are based mainly on Latin language have international
uniformity. By the name Saccharum officinarum, all botanists of the world would understand that
it is sugarcane.
Jones and Luchsinger (1987) have stated that common names present five major problems men-
tioned in the following table:
S. No. Common Names Scientific Names
1. These are not universal. They vary in These are universal and are
different languages. recognised throughout the world.
2. They do not provide information indicating They provide information regarding these
generic and family relationships. relationships.
3. A well-known plant may have hundreds of A well-known plant has only one scientific
common names. name.
4. Sometimes, two or more plants have the Two or more plants always have different
same common name. scientific names.
5. Many species do not have any common names. All known plants have a scientific name.
The scientific name of a plant consists of two separate words. The first word designates the genus
of the organism and the second word designates the species. Details of this two-name system were
first given by Linnaeus (1753) in his famous binomial system of nomenclature.
8.3 BINOMIAL NOMENCLATURE

8.3.1 What is Binomial Nomenclature?
In the earlier days plant names were long and descriptive e.g. in the herbal of Clusius (1583) a spe-
cies of willow is named Salix pumila angustifolia altera. Then, in 1623, Gaspar Bauhin (1560–1624)
devised a plan of adopting two names for each plant in his Pinax Theatri Botanici. But it was Carolus
Linnaeus (1707–1778), the great Swedish naturalist, to whom the actual credit goes for devising and
methodically employing the binomial system of nomenclature. Linnaeus employed this system in the
first edition of his Species Plantarum in 1753. According to this system the scientific name of a plant
consists of two Latin or latinized words: (1) The first is the name of the genus, i.e. generic name or
generic epithet, and (2) the second is the name of the species, i.e. specific epithet. For example, the
80 Plant Taxonomy
botanical name of sugarcane is Saccharum officinarum. The first word (Saccharum) designates the
genus of the plant and the second word (officinarum) designates the species of this genus.
8.3.2 Generic Name

It is always a noun, and written with a capitalised initial letter and the remainder small. The generic
name is also always singular in number. It may have following types of origin:
1. Several generic names may be in honour of the names of well-known persons, e.g. Theophrasta
in honour of Theophrastus and Candollea in honour of A.P. de Candolle.
2. They may be descriptive, with reference to some common characteristics of the included
species, e.g. Cercocarpus (coiled fruit), Xanthoxylum (yellow wood), etc.
3. They may be of poetic or mythological origin, e.g. Theobroma (god’s food).
4. They may also be the aboriginal name of the plants, e.g. Betula and Quercus which were
the old Greek names for Birch and Oak.
8.3.3 Specific Epithet

According to the 1983 recommendations of the International Code of Botanical Nomenclature (Voss,
et al., 1983), all specific epithets be written with a small initial letter. But if the specific epithet is
derived from common names, or from former generic names, or from a person’s name, the initial
letter of the epithet may be a capital letter. In the typed or handwritten matter, both the specific
epithet and generic names should be underlined. They should be printed in italics or boldface. The
authority, written after the specific epithet, is never underlined.
The specific epithet is often an adjective. According to Article 23 of ICBN, a specific epithet
(i) may be a name in honour of a person, or (ii) may be derived from a geographical location, or
(iii) may originate from an old common name, or (iv) may be derived from some characteristics of
the plant, or (v) it may also be named arbitrarily.
8.3.4 Authority
The name of the species is incomplete if it is not followed by full or abbreviated name(s) of the
author(s). For example, Pyrus malus is incomplete. The complete name is Pyrus malus L. where L. is
abbreviated for Linnaeus. Citation of the full or abbreviated form of the author is necessary because
this will verify the date or time of the first valid publication of the name of a particular taxon.
8.4 INTERNATIONAL CODE OF BOTANICAL NOMENCLATURE ICBN

8.4.1 What is ICBN?
The International Code of Botanical Nomenclature (ICBN) is the set of rules and recommendations
dealing with the formal botanical names that are given to plants. Its main aim is that each taxon
or taxonomic group of plants has only one correct botanical name and that is accepted throughout
the world. Two main principles of ICBN are listed below:
(1) Priority is the guiding principle in botanical nomenclature. The ICBN sets the formal starting
date of plant nomenclature at 1 May, 1753, which is the publication date of Species Plantarum
by Linnaeus.
(2) Each botanical name is fixed to a taxon by a type, which is almost invariably dried plant
material usually deposited and preserved in a herbarium.
Very few hard rules of ICBN apply above the taxonomic rank of family. Each new edition of
ICBN supersedes the earlier editions and is retroactive back to 1753.
8.4.2 Who can Change ICBN?

The ICBN can only be changed by an International Botanical Congress (IBC), with the International
Association for Plant Taxonomy providing the supporting infrastructure.
8.4.3 To which Organisms does ICBN Apply?

The ICBN applies not only to plants, as they are now defined, but it also applies to other organ-
isms studied traditionally by botanists, e.g., Cyanobacteria (blue-green algae), fungi, photosynthetic
protists and taxonomically related non-photosynthetic groups. For fossils, there are special provisions
in ICBN.
8.4.4 Whether ICBN is Applicable to Cultivated Plants?

No. For naming of cultivated plants, there is a separate code called the International Code of
Nomenclature for Cultivated Plants (ICNCP).
8.4.5 History of ICBN

Linnaeus in 1737 and again in 1751 proposed the elementary rules of naming plants in his Philosophia
Botanica. Then in 1813, A.P. de Candolle set forth a detailed set of rules regarding plant nomencla-
ture in his Theorie elementaire de la botanique. The same rules of Linnaeus, A.P. de Candolle and
his son Alphonse de Candolle were later evolved into our present International Code of Botanical
Nomenclature (ICBN).
Alphonse de Candolle convened the First International Botanical Congress in 1867 in Paris. It
was attended by the botanists of several countries. They adopted a set of rules of plant nomencla-
ture, most of which were proposed by A. de Candolle. These excellent rules of plant nomenclature
are known as de Candolle rules or Paris Code of 1867. Subsequent meetings of the International
Botanical Congress were held in 1892 (Rochester Code), 1905 (Vienna Code), 1907 (American Code)
and 1910, but a general agreement, regarding the internationally acceptable rules of plant nomencla-
ture, was reached in the meeting of the IBC at Cambridge in 1930. Lawrence (1951) has discussed
the detailed history of the Code and may be quoted that in 1930 at the Cambridge Congress “for
the first time in botanical history, a code of nomenclature came into being that was international in
function as well as in name”. This code is called the International Code of Botanical Nomenclature.
Scientists in the International Botanical Congresses suggest the modifications or amendments which
are incorporated in the ICBN on a regular basis.
82 Plant Taxonomy
The International Code of Botanical Nomenclature, 1983, was adopted by the Thirteenth
International Botanical Congress, Sydney (Australia) in August 1981, and the Chairman of the
editorial committee was E.G. Voss. ICBN is divided into three parts, i.e. Principles, Rules and
Recommendations.
8.4.6 Principles of ICBN

The philosophical basis of the Code is formed by the following six principles:
“1. Botanical nomenclature is independent of zoological nomenclature.
2. The application of names of taxonomic groups is determined by means of nomenclatural
types.
3. The nomenclature of a taxonomic group is based upon priority of publication.
4. Each taxonomic group with a particular circumscription, position, and rank can bear only one
correct name, the earliest that is in accordance with the Rules, except in specific cases.
5. Scientific names of taxonomic groups are treated as Latin regardless of their derivation.
6. The Rules of nomenclature are retroactive unless expressly limited.”
8.4.7 Rules and Recommendations of ICBN

According to the ICBN (1983) the detailed provisions of the Code “are divided into Rules, set out in
Articles, and Recommendations”. As mentioned under point No. 4 of the Preamble of 1983 ICBN
“The object of the Rules is to put the nomenclature of the past into order and to provide for that of
the future; names contrary to a rule cannot be maintained”.
As mentioned under point No. 5 of the Preamble of 1983 ICBN “The Recommendations deal
with subsidiary points, their object being to bring about greater uniformity and clearness, especially
in future nomenclature; names contrary to a recommendation cannot, on that account, be rejected,
but they are not examples to be followed.”
The Rules and Recommendations of ICBN apply to all organisms treated as plants (including
fungi but excluding bacteria), whether fossil or non-fossil. Nomenclature of bacteria is governed by
a separate code called International Code of Nomenclature of Bacteria (ICNB).
It is stated under point No. 10 of ICBN (1983) that the latest “edition of the Code supersedes all
previous editions.”
8.4.8 Some Later Codes

St. Louis Code (1999)
The 16th International Botanical Congress was held at St. Louis, Missouri, USA in August 1999, and
the International Code of Botanical Nomenclature adopted in this Congress is called St. Louis Code.
Botanists of 85 nations attended this Congress. Hectic discussions were made mainly in view of the
facts that “as many as two-thirds of the world’s 3,00,000 plant species are in danger of extinction
in nature during course of the 21st century”, and almost every aspect of the life of human beings
depends on plants. Botanists made some resolutions and recommendations, of which some major
ones are undermentioned:
(A) Resolutions Six resolutions were passed in this International Botanical Congress at St. Louis.
(1) Resolution I Botanists agreed on 10 major aspects in this resolution, of which some are mentioned
below. They call for:
(i) “the establishment of a new coordinating body associated with United Nations to monitor the
status of plants throughout the world, detect those in most danger, and take steps to conserve
them in nature, in botanic gardens, or in gene banks”;
(ii) “securing additional funds for study of plants throughout the world”;
(iii) “making all the information about plants generally available on the Internet”;
(iv) placing additional emphasis on the importance of the “survival of biodiversity throughout the
world”;
(v) maintaining “an active census of the status of each country’s plants” at national level;
(vi) “actively developing and implementing plans to conserve the world’s economic plants ....”;
(vii) “devoting special attention to the conservation of medicinal plants ....”;
(viii) “funding internationally an ongoing programme of research on plant population biology....”
(2) Resolution II Botanists of the 85 nations attending this Congress call on governments and policy
makers to:
(i) “recognize the importance of developing and maintaining scientific expertise, provide
resources for the education and training of scientists, and maintain career opportunities”....
especially in biological sciences;
(ii) “actively develop floras and detailed accounts of plants of all regions ....”;
(iii) “support collaborative programmes between and among developed and developing
countries”;
(iv) “ensure high priority be given to the maintenance of botanical museums, herbaria, libraries,
gardens, living plant collections and gene banks ....”;
(3) Resolution III Botanists in this resolution resolved to:
(i) “increase our knowledge of diversity and relationships of plants ....” and “to make that knowl-
edge accessible to all”;
(ii) “advocate to policy makers the relevance of plant sciences ....”, and thus “maintain the quality
of human life on earth”;
(4) Resolutions IV to VI These three resolutions are regarding the (i) importance and programmes of
biodiversity (Resolution IV); acceptance of the decisions of Nomenclature Section of the Congress
(Resolution V); and selection of sites for future Congresses to facilitate the “attendance of botanists
from all regions of the world” (Resolution VI).
(B) Recommendations Several recommendations have been made in ICBN in the St. Louis
Code by botanists attending the XVI International Botanical Congress, of which only some are
undermentioned:
1. The scientific names under the jurisdiction of the Code, irrespective of rank, “are consistently
printed in italic type. The Code sets no binding standard in this respect, as typography is a
matter of editorial style and tradition, not of nomenclature”.
84 Plant Taxonomy
2. To set off scientific names even better, the use of italics for technical terms and other words
in Latin, “has now been abandoned”.
3. For style of bibliography, the titles of the books are “abbreviated in conformity with Taxonomic
Literature, ed. 2 by Stafleu & Cowan (1976–1988; with supplements by Stafleu & Mennega,
1999–2000)”.
4. For style of bibliography the journal titles are abbreviated in conformity with Botanico-
periodicum-huntianum (1968) and its supplement (1991).
5. Author citations of scientific names are standardized in conformity with Authors of Plant
Names by Brummitt and Powell (1992).
6. The single largest area of change in St. Louis Code concerns typification, where many pro-
posals have been made on Lectotypification (for details refer to original code).
7. All reference to registration of new botanical names, to become “mandatory from a future
date, be deleted from the Code ....”.
8. Fossil plant nomenclature underwent profound changes in St. Louis Code ( for details refer
to original Code).
9. Fungal nomenclature was only affected in a marginal way by decisions of the St. Louis
Congress.
10. One new term “isonym” has been introduced into the Code, “defined to mean the same name
used independently by different authors at different times ....”.
11. In the field of terminology, the terms “homotypic synonym”, “heterotypic synonym” and
“replacement names” were accepted as optional equivalents of the earlier “nomenclatural
synonym”, “taxonomic synonym” and “avowed substitute”.
12. The terminations -viridae, -virales, -virinae, and -virus were “outlawed for names of sub-
classes, orders, subtribes and genera, respectively.
Vienna Code (2005)
The latest XVII International Botanical Congress was held in Vienna in 2005 (XVI being at St.
Louis, Missouri, USA), and the International Code of Botanical Nomenclature accepted in this
Congress is called Viena Code. It is written entirely in English and has been translated in several
other languages. One of the reasons invoked for the choice of Vienna as the site of XVII Congress
was that the second International Botanical Congress had been held there exactly 100 years earlier,
i.e., in 1905. It was in this IInd Congress in 1905 that the first internationally developed rules gov-
erning nomenclature of plants were accepted, and these rules were recognised as Vienna Rules. The
XVII IBC held on 12–16 July, 2005 at Vienna was attended by 198 registered members carrying
402 institutional votes. On the other hand, St. Louis Congress at Missouri, held six years ago in
1999 was attended by 297 members carrying 494 institutional votes.
The Vienna Code does not differ substantially in overall presentation and arrangements from the
St. Louis Code, and the number of Articles remains the same. Some selected additions and recom-
mendations of Vienna Code are listed below:
1. The most notable feature of Vienna Code “is the inclusion for the first time a Glossary, which
appears in Appendix VII. This Glossary is very tightly linked to the wording of the Code,
and “only nomenclatural terms defined in the Code can be included”.
2. The scientific names under the jurisdiction of Code, irrespective of rank, are consistently
printed in italic type. The “Code sets no binding standard in this respect, as typography is
a matter of editorial style and tradition not of nomenclature”.
3. The titles of the books in bibliographic citations are abbreviated in conformity with
“Taxonomic Literature. ed. 2, by Stafleu and Cowan (1976–1988; with 6 supplements by
Stafleu and Mennega, 1992–2000)”, but with capital initial letters.
4. For titles of the journals in bibliographic citations, the abbreviations follow the Botanico-
periodicum-huntianum, ed. 2 (2004).
5. Author citations of scientific names appearing in the Code are “standardized in conformity
with Authors of Plant Names, by Brummitt & Powell (1992)”; these are “also adopted and
updated by the International Plant Names Index”.
6. “Perhaps the most important single decision incorporated into the Vienna Code was to deal
with what many have recognized as a bomb waiting to explode, the publication status of
theses submitted for a higher degree”. The Code decided that “no independent non-serial
publication stated to be a thesis submitted for a higher degree on or after 1 January 1953
would be considered an effectively published work without a statement to that effect or other
internal evidence” (for details, readers should consult the original Code).
7. Regarding valid publication of names, Vienna Code made it clear that “names be composed
only of letters of Latin alphabet, except as otherwise provided in the Code.”
8. St. Louis Code accepted that all fossil taxa should be treated as morphotaxa. In Vienna
Code, this has not been considered appropriate. A distinction between a morphotaxon and a
regular fossil taxon is now established in Vienna Code.
9. In Vienna Code, it was agreed that “the starting date for valid publication of suprageneric
names of spermatophytes, pteridophytes and bryophytes be 4 August 1789, the date of pub-
lication of Jussieu’s Genera Plantarum”.
10. Vienna Code also established that parenthetic author citation is not permitted at suprageneric
ranks.
11. Four family names, previously overlooked in Berchtold and Presl’s rare, later, multivol-
ume work of the same name (1823–1825) have been updated: Aquifoliaceae, Cornaceae,
Potamogetonaceae and Punicaceae.
12. Under the concept of “minimum invalidity” (Art. 33.10), the “rules determining when a rank
is denoted by a misplaced term (and hence not validly published) were clarified and made
more practical”.
13. From 1 January, 2007, a new combination, a new generic name with a basionym, or an allowed
substitute “is not validly published unless its basionym or replaced synonym is cited”.
14. Vienna Code establishes that only if validly published does a name have any status; indeed,
unless otherwise indicated, the word “name” in the Code means a name that has been validly
published”.
86 Plant Taxonomy
8.4.9 Future Botanical Congress and Legality of ICBN

The International Code of Botanical Nomenclature (ICBN) is published under the authority of the
International Botanical Congresses (IBC). The next meeting of the IBC will be held in Melbourne,
Australia, from 23–30 July, 2011. Similar to other international codes of nomenclature, ICBN also
has no legal status and is dependent on the voluntary acceptance of its rules by scientists in general
and botanists in particular.
8.5 SOME IMPORTANT RULES OF NOMENCLATURE

1. Ranks and Endings of Taxa In an accepted system of classification, each individual plant
is treated as belonging to a number of taxa of different ranks. Generally, the species is
considered as the basic unit of classification. Other main ranks in the flowering plants in an
ascending order are genus, family, order, subclass and class. However, ICBN (Voss et al.,
1983) has mentioned 22 different ranks and some standardized grammatical endings (suffixes)
for the ranks from division down to the level of genus. All these ranks are mentioned under
Article No. 1.5 (Table 1.1, Chapter 1).
2. Principle of Priority One plant might have been described under different botanical names
by various plant nomenclaturists in different parts of the world. But, according to the “prin-
ciple of priority” “each taxon is to be known by its earliest name”. For example, Cleome
gynandra Linn. was first described and named by Linnaeus in 1753. Then he himself changed
its name as Cleome pentaphylla Linn. In 1824 de Candolle recognized three separate gen-
era (Cleome, Polanisia and Gynandropsis) and named the Linnean genus as Gynandropsis
pentaphylla (Linn.) DC. Iltis in 1960 merged Gynandropsis and Cleome into one genus,
i.e. Cleome. So according to the “Principle of priority”, the oldest name (Cleome gynandra
Linn.) is the correct valid name.
Because of Principle of Priority, names of several plants have now been changed.
3. Type Method The type method is a legal device to provide the correct name for a taxon. A
type specimen is a herbarium sheet (or rarely a drawing or a photograph) of a specimen which
was used by the author to provide its authentic description. According to Article 9 of ICBN
the type of a genus is a species (e.g. the type of genus Vernonia is V. noveboracensis (L.)
Michx.), and the type of a family is a genus (e.g. Aster is the type genus of Asteraceae).
Several kinds of types designated by ICBN are undermentioned:
i. Holotype It is a specimen used by the author in the original publication as the nomen-
clatural type.
ii. Isotype It is a duplicate specimen of the holotype, i.e. from the same collection, with
the same locality, date and number as the holotype.
iii. Lectotype It is a specimen selected by a competent worker from the original material
studied by the author of the species, when no holotype was designated or when the
holotype has been destroyed or lost.
iv. Neotype It is a specimen selected to serve as a substitute for the holotype when all
material, on which the name of the taxon was based, is missing.
v. Nomenclatural Type It is that element with which the name of a taxon is permanently
associated.
vi. Syntype It is one of the two or more specimens cited by an author of a species when
no holotype was designated; or, a syntype is any one of the two or more specimens
originally designated as types.
vii. Paratype or “Co-types” It is a specimen other than isotype or holotype. If two or more
specimens have been cited as types by the author, the remaining cited specimens are
called “Co-types” or paratypes.
viii. Topotype It is a specimen collected from the same locality from where the holotype
was collected.
4. Synonyms and Related Definitions A name rejected due to misuse or difference in taxo-
nomic judgement is called synonym.
A specific or intraspecific name which has priority and is retained when transferred to a
new taxon, is called a basionym.
A case in which two or more identical names are based on different types, of which only
one can be a legitimate name, is called a homonym.
An illegitimate binomial, in which the name of the genus and the name of the species is the
same, is called a tautonym, e.g. Armoracia armoracia (L.) Britton.
An automatically created legitimate tautonym for infraspecific or infrageneric taxa is called
an autonym, e.g. Hypericum subgenus Hypericum section Hypericum.
5. Citation of Author Some of the rules related to author citation are undermentioned:
i. Original Author The name of a taxon is complete and accurate only when it is followed
by a full or abbreviated form of the author(s) who first validly published the concerned
name, e.g. Liliaceae and Lilium superbum are incomplete; the complete names are
Liliaceae Adans. and Lilium superbum Linn.
ii. Joint Author If two authors have jointly published the name of a taxon, the names of
both the authors should be cited and linked by the words et or & e.g. Illicium griffithi
Hook & Thoms (or Hook et Thoms).
iii. Rank Alteration When a taxon of a lower rank is upgraded in a higher rank but retains
its name, the author’s name who published it first should be cited in the bracket It is to
be followed by the name of the author who made the alteration, e.g. Allioni raised the
rank of variety Medicago polymorpha var. orbicularis L. to the species rank. Therefore,
it becomes Medicago orbicularis (L.) All.
iv. Name Proposal When the name of a taxon is proposed but not validly published by
one author, and is later on validly published by another, the word ex should be used as a
connecting link between the name of the former author and the name of the subsequent
author, e.g. Gossypium tomentosum Nutt ex Seem.
6. Names of Cultivated Plants Wild plants brought under cultivation retain their original
names.
88 Plant Taxonomy
7. Latin Diagnosis The diagnosis, details and description of new taxa published before 1st
January, 1955 were accepted by ICBN as valid, irrespective of the language. After this date
the description of any new taxa would be considered valid only if accompanied by Latin
diagnosis.
8. Effecttve and Valid Publication The publication of new names and description are effective
and considered valid only when they are distributed in a printed form to the general public
or to at least ten well-established botanical institutions.
A validly published names, as specified in the ICBN, is the one in which the basic provi-
sions are (1) effective publication, (2) publication in the form specified for the name of each
category of taxa, (3) publication with a description, or a reference to a previously published
description, of the taxon to which the name applies, (4) accompanied by a Latin description
or by a reference to a previously and effectively published Latin description of the taxon,
and (5) an indication of the nomenclatural type. On the other hand, an effectively published
name is the one published in printed matter generally available to botanists.
9. Choice of Names when the Taxon Rank is Changed When the rank of a taxon is changed
(viz. a species becomes a genus or vice-versa) the earliest legitimate name in its new rank
is its correct name.
10. Choice of Names when same-rank Taxa are United When two or more taxa of the same
rank (viz. two or more genera, two or more species, etc.) are united into one, the oldest
legitimate name of these taxa would be retained as the name of the united taxon.
11. Retention of Names of Divided Taxa When a genus or a species is divided into two or more
genera or species, respectively, the original name of the genus or species must be retained.
12. Retention of Names of Taxa on Transference When a subdivision of a genus is transferred
to another genus, or a species is transferred to another genus without the change of the
rank, the original legitimate name must be retained. For example, Hydrocotyle asiatica L.,
on transference to genus Centella, must be named as Centella asiatica (L.) Urban.
13. Rejection of Names Names of taxa must be rejected in the following conditions:
i. When the names are illegitimate (i.e. if it is a tautonym, later homonym, rejectable generic
name, nomenclaturally superfluous, etc.).
ii. When the names give different meanings, and become a permanent source of
confusion.
iii. When the characters of the name of the taxon are derived from two or more different
elements.
iv. When the generic names coincide with a morphological term, or are unitary designations
to species, or are words not intended as names.
v. When the specific names are tautonyms, or are published without any consideration of
binomial system, or are words not intended as names.
14. Names of Different Taxa According to the International Code of Botanical Nomenclature
(1983) the following should be the pattern of naming different taxa:
i. Genera and higher ranks should be monomials, e.g. Aesculus L., Rosa L., Rosaceae
Juss.
ii. Species should be binomials, e.g. Gossypium tomentosum Nutt.

iii. Subspecies should be trinomials, e.g. Hibiscus moscheutos ssp. palustris (L.) Clausen.
iv. Varieties should be quadrinomials, e.g. Lilium catesbaei Walter ssp. catesbaei var. longii
Fernald.
B BOTANICAL NAMES
Botanical names are either Latin words or words that have been latinized from Greek or some other
language.
The name of a species is a binomial and consists of a generic name and a specific epithet. A
specific epithet is the second part of the binomial. However, one must be quite clear about the fact
that it is wrong to use the specific epithet alone to designate a particular species; it must always be
used with a generic name to form the binary combination for that species. The specific epithets are
formed from nouns, adjectives, etc. and may join these words with a large number of prefixes and
suffixes.
8.6 COMMON PREFIXES USED IN SPECIFIC EPITHETS

Some Latin prefixes of numbers are as follows:
1. uni-(L.): uniflorus (one-flowered)
2. bi-(L.): bifoliatus (two-leaved)
3. tri-(L.): triangularis (with 3 angles)
4. quadri-(L.): quadrangularis (with 4 angles)
5. quinque-(L.): quinquefolius (5-leaved)
6. sex-(L.): sexangularis (6-angled)
7. septem-(L.): septemlobus (7-lobed)
8. octo-(L.): octoflorus (8-flowered)
9. noveme-(L.): novemneris (with 9 nerves)
10. decum-(L.): decumlobus (with 10 lobes)
Some of the Greek prefixes of numbers include mon- (for one, e.g. monandra), di- (for 2), tri-
(for 3), tetra- (for 4), penta- (for 5), hex- (for 6), hepta- (for 7), octo- (for 8), ennea (for 9) and
deca- (for 10, e.g. decapetalus).
Some of the other common prefixes of numbers are amphi- (Gr.): for two kinds; diplo- (Gr.): for
double; haplo- (Gr.): for single; multi- (L.): for many; poly-(Gr.): for many; a- or ab- (L.): away
from; ecto- (Gr.): outside; endo- (Gr.): inside; inter- (L.): between, and; intra- (L.): within.
8.7 COMMON SUFFIXES USED IN SPECIFIC EPITHETS

Some of the common suffixes along with one example of each of them are undermentioned:
-aceus: crustaceus; -alis: digitalis; -aris: angularis; -arium: aquarium; -aticus: aquaticus; -estris:
campestris; -eus: roseus; -ilis: sexitilis; -osus: foliosus.
90 Plant Taxonomy
8.8 PLANT PARTS USED AS EPITHETS

Almost all plant parts, including root, stem, leaf, bud, flower, fruit and seeds, have been used as
names of specific epithets. Some of the specific epithets, along with the related plant parts in paren-
thesis, are undermentioned:
receptaculum (for receptacle), sepalum (for sepals), carpellum (for carpel), loculus (for locules), stylus
(for style), petalum (for petals), discus (for disc), ovarium (for ovary), ovulum (for ovule), pedicellus
(for pedicel), pistillum (for pistil), fructus (for fruit), sperma (for seed), folium and phyllon (for leaf),
rhiza (for root), caulos (for stem), etc.
8.9 SPECIFIC EPITHETS LINKED WITH COLOUR

Some of the specific epithets linked with colour, along with the name of the colour in the parenthesis,
are undermentioned alphabetically:
albicans (whitish), albus (white), atrovirens (dark green), aureus (golden yellow), calcareus (chalky
white), candidus (shining white), croceus (saffron yellow), flavidus (slightly yellow), flavus (pale
yellow), fulvus (dull yellow), glaucus (grey-green), luteolus (pale yellow), niger (black), niveus (snow
white), purpureus (purple), roseus (rosy), violaceous (violet), virens (green) and, viridis (green).
8.10 SPECIFIC EPITHETS LINKED WITH GEOGRAPHY

Some of the geography-linked specific epithets, along with their related geographical places in
parenthesis, are undermentioned alphabetically:
africanus (of Africa), americanus (of America), arabicus (of Arabia), argentinus (of Argentina),
asiaticus (of Asia), australiensis (of Australia), austriacus (of Austria), brasiliensis (of Brazil),
canadensis (of Canada), chinensis (of China), cubensis (of Cuba), europaeus (of Europe), germani-
cus (of Germany), helveticus (of Switzerland), indicus (of India), italicus (of Italy), mexicanus (of
Mexico), sinensis (of China), virgnicus (of Virginia) and, zeylanicus (of Ceylon).
8.11 SPECIFIC EPITHETS LINKED WITH SIZE

Some of size-linked epithets are mentioned below alphabetically:
altus (altitude or tall), exaltatus (very tall), giganteus (gigantic or very large), grandis (large), humilis
(dwarf), major (greater), minor (less), minutus (very small), procerus (very tall), pumilus (dwarf)
and, robustus (stout or robust).
8.12 SPECIFIC EPITHETS LINKED WITH HABIT

Epithets, along with the name of the related habit in parenthesis, are undermentioned
alphabetically:
arborescens (arborescent), dichotomus (dichotomous), erectus (erect), furcatus (forked), prostratus
(prostrate), ramosus (branched), repens (creeping) and, stoloniferus (stoloniferous).
8.13 SPECIFIC EPITHETS LINKED WITH HABITATS

Some of such epithets, along with the name of the related habitat in the parenthesis, are listed below
alphabetically:
amphibius (living both on land and in water), aquaticus (living in water), arvensis (growing in
ploughed field), campestris (of field), hypogeus (underground), lacustris (of ponds or lakes), litto-
ralis (growing on seashores), maritimus (of sea), montanus (of mountains), palustris (of swamps),
rupestris (on rocks), sativus (cultivated), sylvaticus (of woods), sylvestris (growing in woods) and,
terrestris (growing in dry soil).
C PHYLOCODE: A NEW SYSTEM OF NOMENCLATURE
8.14 WHAT IS PHYLOCODE?

Recently, Cantino and de Queiroz (2001) proposed a new code for naming organisms by clear “ref-
erence to phylogeny (ancestry and descent), rather than on the basis of the Linnaean hierarchy of
taxonomic categories (species, genus, family, and so on)”, as stated by Robinson and Kommedahl
(2002), and this “new code” has been named by then as “Phylocode”. They also proposed a new
term “clades” in this code. According to them, “clades” is a “group(s) of species comprising a com-
mon ancestor and all its descendants”. Clades are the constituents of the “tree of life”. All clades
should have explicit and unambiguous names that do not change with time. The current systems of
nomenclature are based on Linnaean hierarchy, and they do not “boast such immutability”.
The International Code of Phylogenetic Nomenclature is known in short as “Phylocode”. It is a
developing “draft for a formal set of rules governing phylogenetic nomenclature”. The current version
of Phylocode is specifically designed “to regulate the naming of clades, leaving the governance of
species names up to the rank-based codes”.
8.14.1 How has Phylocode Come into Existence?

Phylocode is not a new idea. It is actually “based on ideas presented in the literature since the late
1980s and, more formally, on the outcome of a workshop held at Harvard University in August 1998”
(Robinson and Kommedahl, 2002). In this workshop, a draft proposal was put forward, and this draft
covers only the naming of clades. It was also decided in this workshop that “rules governing species
names will be added later”. However, as a temporary measure, “Linnaean binomial nomenclature is
used in the draft Phylocode where species names are needed”.
8.14.2 How are the Species Named in Phylocode?

As mentioned above under Article 8.14.1, the final rules governing species names have not been
decided yet. According to Robinson and Kommedahl (2002), “the form that species names should
take in the Phylocode” is still controversial. Cantino et al. (1999) earlier presented 13 possibilities
for naming species phylogenetically and compared these methods with each other and also with
the Linnaean system”. It is, however, for the scientific community to finally decide whether or
92 Plant Taxonomy
not the “Phylocode should become the sole code governing the names of the taxa” (Robinson and
Kommendahl, 2002).
8.14.3 What is the Basic Difference between Phylocode and Linnaean

Binomial Nomenclature?
The fundamental characteristic “that distinguishes the Phylocode from the conventional hierarchic
nomenclatural systems is its ranklessness” according to Robinson and Kommendahl (2002). The
proposed Phylocode will cover the naming of clades and species, but in this Code, these terms
will refer not to ranks but to different kinds of biological entities. It is so because in Phylocode
“both (clades and species) are the products of evolution that are discovered, rather than created, by
systematists, and both have an objective existence regardless of whether they are named” (Cantino
and de Queiroz, 2001).
8.14.4 What would be the Starting Date for Phylocode?

Regarding the starting date for Phylocode, Robinson and Kommedahl (2002) stated that the “start-
ing date for the new Code, which has not yet been decided, will coincide with the publication of a
companion volume providing definitions of widely used clade names”.
8.14.5 What does Phylocode Advisory Group now want from the
Scientific Community?
There is an advisory group which is now working on formulation and implementation of Phylocode.
This group is now coordinating work and seeking comments and ideas concerning this proposal from
as many people as possible. Anybody interested is welcome to review the current draft of Phylocode
and communicate it to the Phylocode Advisory Committee.
8.15 PRINCIPLES OF THE PHYLOCODE

The proposed Phylocode is based on certain principles. It allows freedom of taxonomic opinion
with reference to hypotheses about relationships. It only deals how the names are to be applied with
reference to a given phylogenetic hypothesis. As published in Division 1 (Principles) of Phylocode,
the five basic principles of Phylocode are reference, clarity, uniqueness, stability, and phylogenetic
context.
(1) Reference The most basic purpose of names of taxon is to “provide a means of referring to
taxa, as opposed to indicating their characters, relationships, or membership”.
(2) Clarity The names of taxon should be “unambiguous in their designation of particular taxa.
Clarity in nomenclature is achieved through explicit definitions.
(3) Uniqueness For promoting clarity, “each taxon should have only one accepted name, and each
accepted name should refer to only one taxon”.
(4) Stability During course of time, the names of taxa should not change. “As a corollary, it must
be possible to name newly discovered taxa without changing the names of previously discovered
taxa”. There should, therefore, be a complete stability in the names of taxa.
(5) Phylogenetic context The concern of Phylocode is only with the “naming of taxa and the
application of taxon within a phylogenetic context”.
8.16 PHYLOCODE: AN OVERVIEW

The phylogenetic nomenclature will be regulated by the Phylocode because the latter will provide
rules for the following:
(1) How to decide which combinations of names and definitions will be considered validly pub-
lished (for details, refer to Chapter II of Phylocode)?
(2) Which of the names and definitions will be considered homonyms or synonyms (for details,
refer to Articles 13 and 14)?
(3) Which one of a set of homonyms or synonyms will be finally considered vaild?
Furthermore, the Phylocode will only allow:
(i) the naming of clades (for details, refer to Article 1.1), and
(ii) the use of specimens, species, and amorphies as specifiers ( for details, refer to
Article 11).
8.17 REGISTRATION DATABASE OF PHYLOCODE

8.17.1 RegNum
The future of the Phylocode is still undecided. If and when it is finally implemented, the “Phylocode
will be associated with a registration database called RegNum. The RegNum will store the names
of all clades and definitions that will be considered potentially valid”.
The utility of RegNum will be to “provide a publicly-usable tool for associating clade names with
definitions”. In the latter stages, this will be “associated with sets of subtaxa or specimens through
phylogenetic tree databases (such as TreeBASE)”. RegNum will also be an important tool in taking
a decision that “which one of a number of synonyms or homonyms will be considered valid”.
8.18 MAIN EVENTS IN THE HISTORY OF PHYLOCODE

1. As mentioned earlier under Article 8.14.1, the idea of Phylocode came as an outcome of a
workshop of systematists held at Harvard University in August 1998. An advisory group of
leading systematists was formed in this workshop.
2. In April 2000, these systematists prepared a draft of Phylocode and made it public on the
web. They also invited comments from the systematists globally.
3. In July 2002, a second workshop on Phylocode was held at Yale University, USA. Several mod-
ifications were made in the rules and recommendations of Phylocode in this workshop.
4. From July 6, 2004 to July 9, 2004, the First International Phylogenetic Nomenclature Meeting
took place in Paris (France). It was attended by about 70 systematists and evolutionary biolo-
gists from 11 nations. A society was formed, namely International Society for Phylogenetic
Nomenclature (ISPN). An advisory group was formed to have a complete control on the
development of Phylocode.
94 Plant Taxonomy
5. From June 28, 2006 to July 2, 2006 the Second International Phylogenetic Nomenclature
meeting took place at Yale University, USA.
6. From July 21, 2008 to July 23, 2008, the Third International Phylogenetic Nomenclature
meeting was held at Dalhousie University, Halifax, Nova Scotia, Canada.
8.19 FUTURE OF PHYLOCODE

Still there exists some definite criticism of the Phylocode, and therefore, the Code as such is still
controversial. Till the late 2007, the “number of supporters for the official adoption of Phylocode is
still small”, and it is still uncertain when, actually, “the Code will be implemented, and how widely it
will be followed”. Majority of its supporters believe that the registration database or RegNum should
be made popularised as much as possible among the scientific community, as a first major step in
this direction. A lot, however, depends on following the recommendations of the Third International
Phylogenetic Nomeclature meeting held at Dalhousie University, Canada between July 21, 2008 and
July 23, 2008.

1. Instead of common names, why do we need scientific names for all plants? Give three dif-
ferences between common names and scientific names.
2. What is binomial nomenclature? Who proposed it and when?
3. What is International Code of Botanical Nomenclature? It applies to which organisms? Who
can make changes in ICBN?
4. Write any four principles of ICBN.
5. Give an account of resolutions and recommendations of St. Louis Code.
6. Write at least six major recommendations made by Vienna Code.
7. The XVIII meeting of International Botanical Congress is scheduled to be held in July, 2011.
Name the host city and country of this meeting.
8. Define: (a) holotype, (b) isotype, (c) lectotype, and (d) neotype.
9. What is the difference between a validly published name and an effectively published
name?
10. Give a brief account of phylocode: a new system of nomenclature.
Suggested Reading
Cantino, P.D. and de Queiroz K., 2001, Phylocode: a Phylogenetic Code of Biological Nomenclature (draft
document): www.ohiou.edu/phylocode/printable.html.
________, Bryant, H.N. and Lee M.S.Y., 1999, Species names in phylogenetic nomenclature. Syst. Biol.
48: 790–807.
Clausen, R.T., 1938, On the citation of ‘authorities’ for botanical names, Science 88: 299–300.
________ 1941, On the use of the terms “subspecies” and “variety”, Rhodora 43: 157–167.
Fosberg, F.R., 1942, Subspecies and variety, Rhodora 44: 153–157.
Greuter, W. (ed.), 2000, International Code of Botanical Nomenclature. Regnum Vegetabile. 138: Konigstein
(Germany), 16th Int. Bot. Cong. (St. Louis Code).
Heller, J.C., 1964, The early history of “binomial nomenclature”, Huntia 1: 33–70.
Jeffrey, C., 1973, Biological Nomenclature, Edward Arnold Ltd., London.
Johnson, A.T., 1971, Plant Names Simplified, W.H. & L. Collingeridge Ltd., London.
Lawrence, G.H.M., 1951, Taxonomy of Vascular Plants, Macmillan, New York.
McVaugh, R., R. Ross and F.A. Stafleu, 1968, An Annotated Glossary of Botanical Nomenclature, Regnum
Vegetabile, Bohn.
Nicolson, D.H., 1974, Orthography of names and epithets: Latinization of personal names, Taxon 23:
549–561.
________ and R.A. Brooks, 1974, Orthography of names and epithets: Stems and compound words, Taxon
23: 163–177.
Radford, A. E., 1986, Fundamentals of Plant Systematics, Harper & Row, New York.
Robinson, P. and T. Kommedahl, 2002. Phylocode: A New System of Nomenclature, Sci. Editor 25(2):
52.
St. John, H., 1958, Nomenclature of Plants, Ronald, New York.
Smith, A.C., 1957, Fifty years of botanical nomenclature, Brittonia. 9: 2–8.
Voss, E.G. (ed.), 1983, International Code of Botanical Nomenclature (ICBN), Regnum Vegetabile, Vol.
97. Bohn, Scheltema & Holkema, Utrecht.
C
H
A
P
T
E
R
MODERN TRENDS
IN PLANT
TAXONOMY 9
It is true that morphological characters of the plants have provided the foundation and framework for
taxonomy. These characters have been used extensively in the preparation of classification systems,
diagnostic keys, etc., and are still indispensable to the systematists. But in the light of the modern
developments, the complete knowledge of taxonomy is possible only when a synthetic approach, in
the light of the principles of various disciplines, is applied. These disciplines (viz. plant anatomy,
cytology, palynology, embryology, physiology, ecology, genetics, plant geography, etc.) have played a
significant role in plant taxonomy. The old ideas of classification and phylogeny of the plants have
changed enormously because of the newer aspects of taxonomy such as biosystematics, cytotaxonomy,
chemotaxonomy, molecular taxonomy, etc.
Applicability of the data of some of these branches (anatomy, cytology, palynology, etc.) of botany
to the modern taxonomic procedures are discussed briefly in this chapter.
9.1 EXTERNAL MORPHOLOGY IN RELATION TO TAXONOMY

According to Radford (1986) the morphological data of plants are easily observable and obtain-
able, and are “thus used most frequently in taxonomic studies”. The evidence from the external
morphology provides the “basic language for plant characterization, identification, classification and
relationships” (Radford, 1986).
Angiosperms are identified by morphological characters. Natural groups are defined mainly with
the help of floral characters. The basic similarity in the morphology of flowers, fruits and seeds
within different species, genera, families and orders provides a sound base in characterising taxo-
nomic groups. Morphology provides majority of the characters used in the construction of systems
of taxonomy. Therefore, morphology is closely related to taxonomy and would continue to reign over
it for many more years to come (Ogura, 1964; Holttum, 1968; Cronquist, 1975).
Some of the vegetative characters that play a major role in plant taxonomy and in deducing phy-
logeny include growth habit, phenological characters, underground organs, stem, leaves, petiole and
stipules.
Modern Trends in Plant Taxonomy 97
The floral characters which are used commonly in plant taxonomy include types of inflorescence
and flower, perianth structure, floral symmetry, union of floral leaves in each whorl, types of androe-
cium and stamens, gynoecium and carpels, ovules, and also the characters of bracts, bracteoles and
pedicels. Types of fruit and seed also provide good diagnostic features useful at various levels of
classification.
The specific examples showing such a relationship are mentioned below:
1. Growth habit (herbs, shrubs, and trees) may be variable or constant within a genus or a family,
e.g. all Brassicaceae members are herbaceous. Members of Asteraceae are both herbaceous
as well as woody.
2. Growth patterns have been used in defining taxonomic groups above the generic level in
some families, e.g Zingiberaceae.
3. Seedling characters, such as germination, cotyledonary characters, leaf dimorphism, etc. have
been of systematic value in Convolvulaceae (Sampathkumar, 1982), Brassicaceae (Gomez–
Campo and Maria, 1974), etc.
4. Underground parts, such as roots, tubers, etc. are of some taxonomic value in plants. Tubers
are helpful in the taxonomy of Dioscorea and Cyperaceae, whereas in Ranunculus and
Aristolochia the species are delimited on the basis of the shape of the root.
5. Leaf characters, such as arrangement, type, form, duration and venation are widely used in
both the classification and identification. In Ulmus and Betula, the species are delimited only
on the basis of leaf characters. In Trifolium, the species are separated on the basis of stipule
morphology. Dalbergia species are distinguished on the basis of their leaflet size, shape and
arrangement on the rachis. Various venation patterns have been described in the leaves of
Glossopteris by Pant (1958).
6. Inflorescence and flower are the two main organs on which the classification systems and
several other principles of taxonomy are based. Species are distinguished on the basis of calyx
types in Anthyllis, and shape and degree of inflation of bracts in Calystegia. Nectaries and
floral discs are of great diagnostic value in Brassicaceae. Species are distinguished on the
basis of staminodes in Scrophularia, anthers in Eucalyptus, staminal appendages in Alyssum
and degree of branching of inflorescence in Nepetia.
Pollen wall ornamentations have yielded very useful taxonomic information.
A detailed account of the floral characters and their impact on taxonomy has been given by
Lawrence (1951) and Davis and Heywood (1963).
7. Fruit characters are used in distinguishing different families of Rhoeadales. Genera are
delimited in Moraceae on the basis of fruit characters.
8. Seed characters have been used widely in the construction of diagnostic keys and in dis-
tinguishing species. The generic separation of Glinus Linn. and Mollugo Linn. is based
mainly on seed characters, the former bears appendaged seeds while the latter bears non-
appendaged seeds. Features of seeds such as colour, shape and sculpturing prove helpful in
distinguishing species in Drymaria of Caryophyllaceae. The number and shape of the seeds
is the only reliable distinction between Anthericum and Chlorophytum of Liliaceae. Two
genera of Cruciferae (Sisymbrella and Nasturtium) are distinguished on the basis of the
98 Plant Taxonomy
presence or absence of mucilage on testa of seeds. Several species and genera of Acanthaceae,
Asclepiadaceae, Convolvulaceae and Malvaceae are distinguished on the basis of their seed
characters.
9.2 VEGETATIVE ANATOMY IN RELATION TO TAXONOMY

In determining relationship between different genera, families, orders and other taxonomic categories,
the anatomical characters are most useful. Anatomical data have also solved several phylogenetic
problems. Metcalfe (1968) identified several herbarium specimens using vegetative anatomy.
According to Radford (1986) some of the basic evidentiary anatomical characters of well-estab-
lished taxonomic value are the type, size, shape, wall sculpture and pattern of wood cells, stelar
patterns, types of vascular bundles, rays, ground tissue and parenchyma, epidermal and mesophyll
tissue, stomata, trichomes, sclereids, nodes, phloem cells, etc. A few of them are discussed below.
9.2.1 Epidermis
Shape, wall thickness, wall sculpturing and inclusions in the epidermal cells are some of the epi-
dermal characters of taxonomic importance in different families and genera. Occlusion of stomata
by an unidentified substance in Winteraceae, papillate epidermal cells in Graminae, presence and
distribution of silica bodies in Cyperaceae, sclerification of the wall of the epidermal cells in some
genera of Compositae, and presence of very narrow epidermal cells in Stylidiaceae are some of the
characters of useful taxonomic importance.
Sharma and Shiam (1984) described the taxonomic importance of silica bodies in 22 Indian spe-
cies of Cyperus. They also reported the presence of cuticular papillae as a character of taxonomic
significance in Cyperus pilosus, and formation of crater-like pores in C. flabelliformis (Sharma and
Shiam, 1981 a, b).
9.2.2 Stomata
Studies on the morphology and development of different stomatal types are expected to provide clues
to various evolutionary trends among families of angiosperms. They may also prove to be helpful
in assigning taxa of uncertain affinities to proper positions (Paliwal and Anand, 1978). Some of
the stomatal characters of taxonomic significance are their morphology and ontogeny, number and
arrangement of subsidiary cells, and their relationship with other epidermal cells.
About 31 different patterns of stomata and subsidiary cells have so far been reported in vascular
plants by different workers, including Pant (1965), Stace (1965) and Van Cotthem (1970). These
stomatal patterns appear “to be most valuable at higher taxonomic levels” (Jones and Luchsinger,
1987), e.g. these patterns have been used to characterise the subclasses of monocotyledons.
Significance of stomata has been confirmed in the taxonomy of Gramineae (Prat, 1932),
Epacridaceae (Watson, 1962), Combretaceae (Stace, 1965), Rubiaceae (Pant and Mehra, 1965),
Acanthaceae (Paliwal, 1966), Umbelliferae (Guyot, 1971), Papilionaceae (Kothari and Shah, 1975),
Araliaceae and Umbelliferae (Kannabiran and Krishnamurthy, 1979), Myrtaceae (Vauwyk et al.
1982), Cyperaceae (Sharma and Shiam, 1984), and several other angiospermic families.
9.2.3 Trichomes
Epidermal hairs or trichomes are of much taxonomic significance because they exhibit great diversity
of form, size, structure and function. Certain species of Vernonia are differentiated on the basis of
their trichomes. They are also of great value in analysing the suspected hybrids of Compositae and
several other families.
Trichome types and their distribution are useful characters in distinguishing various genera of
Fabaceae and Icacinaceae. Position of Nyctanthes in Oleaceae has been confirmed by Inamdar (1967)
on the basis of the structure and ontogeny of trichomes. Presence of sessile glandular hairs in both
Typhaceae and Sparganiaceae (Solereder and Meyr, 1933) confirms their close affinity with each
other. Cruciferae has been divided into tribes and genera mainly on the basis of types of trichomes
(Schulz, 1936). Species of Digitalis have been divided into two groups mainly on the basis of pres-
ence or absence of glandular hairs. Trichomes in Parthenium argentatum (Compositae) are T-shaped
while in P. incanum they are whiplike with a long thread. Species in various genera of Labiatae are
also separated on the basis of presence or absence of glandular hairs. Trichomes have been the main
basis of the formation of generic key for the Indian members of Compositae (Ramayya, 1969).
9.2.4 Stem Anatomy

Anatomy of vegetative structures has been of importance in separating higher categories, such as
gymnosperms from angiosperms, and monocots from dicots, and not commonly so at lower levels.
But, on the basis of stem anatomy, Oleaceous genus Nyctanthes is considered to have a closer affin-
ity to Verbenaceae (Stant, 1952). In the subgenus Genuini of Juncus, Stace (1970) has shown that
anatomy of the stem can be used to distinguish the majority of its British species. Dioscorea species
are also distinguished on the basis of stem anatomy.
Transformation of cortex into transfusion tissue in Casuarina, structure of stem endodermis in
families such as Asteraceae and Piperaceae, presence of bicollateral vascular bundles in two alternate
rings in Cucurbitaceae, and occurrence of cortical and medullary bundles in some families such as
Amaranthaceae, Chenopodiaceae and Nyctaginaceae, are some of the features of taxonomic impor-
tance. For a detailed study, readers may refer to the profusely illustrated two volumes of Metcalfe
and Chalk (1950).
9.2.5 Nodal Anatomy

According to Dickison (1975) correlations of nodal anatomy with some other features might help
significantly in tracing the phylogeny of angiosperms. Philipson and Philipson (1968) also attempted
to use nodal anatomy as an aid to taxonomy.
Sinnott (1914) considered the trilacunar node as primitive, and unilacunar and multilacunar
nodes as advanced. In all members of Centrospermales, the node is unilacunar. However, there may
be variations in the nodal anatomy even within a single plant. Subfamily Icacinoideae of family
Icacinaceae may be divided into two sections, one with unilacunar nodes and the other characterized
by trilacunar nodes.
According to Paliwal and Anand (1978) majority of dicotyledons possess trilacunar nodes.
Unilacunar nodes are found in Laurales, Caryophyllales, Ericales, Ebenales, Primulales, Myrtales
100 Plant Taxonomy
and a few Tubiflorae and Asteridae members. According to them, multilacunar nodes are found in
Magnoliales, Piperales, Trochodendrales, Umbellales and Asterales.
9.2.6 Petiole Anatomy

Metcalfe and Chalk (1950) and Howard (1963) have suggested that the petiole anatomy might also
be of taxonomic significance. According to Howard (1963) families, genera and even species in some
cases may be identified by petiole characters, such as its position on stem, presence or absence of
stipules, its vascularization, nodal structure, number of traces, etc.
Petiole anatomy of 64 species of Baphia of Leguminosae (Soladoye, 1982) and some species of
Phlomis and Eremostachys of Labiatae (Azizian and Cutler, 1982) provide clear support of its use
in the taxonomy of these genera. Petiolar vascularization has also been helpful in the classification
of Rhododendron and some other genera.
9.2.7 Leaf Anatomy

Leaf anatomy provides various characters of taxonomic importance. Leaf anatomical studies associ-
ated with C4- photosynthetic pathway have resulted (Brown, 1975) in a revised classification of several
genera of the grass family. A prominent chlorenchymatous bundle sheath is present in C4- plants.
Characters of taxonomic significance in leaf anatomy include nature and thickness of epidermis,
mesophyll types, pattern of sclerenchyma, venation patterns, crystals, etc.
Koyama (1967) and Govindrajalu (1966, 1968, 1974, 1975 and 1976) studied the leaf anatomy of
several species of Cyperaceae and formulated keys to identify various species of Cyperus, Fuirena,
etc.
Leaf anatomy has been used widely in several taxonomically different groups such as Euphorbiaceae,
Cyperaceae and Gramineae of Angiosperms and Coniferae of Gymnosperms. Benson (1962) in
Ranunculus, Webster (1967) in Euphorbia and Chamaesyce, Metcalfe (1968) in several genera of
Cyperaceae, and Vidakovie (1957) in Pinus have used several characters of leaf anatomy in differ-
entiating species.
Patterns of the distribution of sclerenchyma in Carex and Festuca have been used in distinguish-
ing species. Sclerenchyma is also used in differentiating two genera of Velloziaceae viz. Vellozia and
Barbacenia. Taxonomic implication of leaf anatomy of several genera of Musaceae, Zingiberaceae,
Xanthorrhoeaceae and Ericaceae has also been established by several workers.
9.2.8 Sclereids
Sclereids are the cells with very thick lignified walls. They have been used as the diagnostic tools in
several taxa. They are extremely rare in monocots, except in certain genera of Araceae, Agavaceae,
Arecaceae and a few other families. In dicots, they are more common in woody forms than in
herbaceous ones. Rao and Das (1981) have shown their taxonomic value in about 30 species of
Limonium.
The systematic value of sclereids has been specifically ascertained in Limoniaceae, Nymphaeaceae,
Theaceae, Oleaceae, Connaraceae, and a few genera of Araceae, Acanthaceae, Ericaceae and
Melastomaceae. The forms of sclereids may be characteristic of a species or genus, and may also
be of taxonomic value.
9.2.9 Specialised Cells and Cell Contents

Specialised cells of peculiar morphology and function are also important diagnostic tools in many
taxa. Motor cells are characteristic of grass leaf epidermis while the bulliform cells occur in the
epidermis of many monocotyledonous families. Silica cells are common in sedges while the presence
of latex-containing cells is the diagnostic feature of succulent plants.
Microscopic characters of cell contents such as starch grains (e.g. Solanum tuberosum), protein
bodies (some Cactaceae), albuminoids (e.g. Laportea), large silica bodies (Musaceae, Arecaceae
and Zingiberaceae), calcium oxalate crystals (e.g. Eichhornia, Allium), cystoliths (e.g. Moraceae
and Urticaceae), and tanniniferous cells (e.g. Xyridaceae), etc. are important diagnostic tools, and
at times prove extremely helpful in delineating species, genera and families.
According to Nair et al. (1977) the epidermal cells of the male plants of Myristicta fragrans
contain simple crystals while those of the female plants contain compound crystals or druses.
9.2.10 Wood Anatomy

The characters of secondary wood are the most important anatomical features that have been used
in resolving problems of taxonomy and phylogeny. “The evolutionary series of vessel elements has
been used in combination with other morphological features to develop hypotheses about phylogeny
of angiosperms” (Jones and Luchsinger, 1987). Bailey (1957), Swamy (1958), Stern (1978), Gregory
(1980), Purkayastha (1980) and Fahn (1987) have clearly established the role of wood anatomy in
taxonomy.
The secondary xylem of different species varies in the size, shape, arrangement, and relative
amounts of its constituents. These variations have been the major source of development of keys by
which plant species can be identified. Botanical species have also been identified from a piece of
timber by observing its anatomical features. Such an identification is of commercial and industrial
importance. Clarke (1938), Phillips (1948) and Fahn (1987) have prepared keys for differentiating
several genera and species of gymnosperms and angiosperms by using characters of wood anatomy.
Keys have also been prepared for separating species of Frankenia, Phyllyrea and Marrubium by
using wood anatomy characters.
Wood anatomy has prompted the allocation of Amborella, Tetracentron and Trochodendron to
their respective independent families. It has also played its role in the institution of a new taxon, the
Degeneriaceae (Baily and Smith, 1942); retention of Sarcandra in Chloranthaceae (Swamy, 1953),
and transfer of Idenburgia from Monimiaceae and its inclusion in Nouhuysia.
Wood anatomy has been used at almost all taxonomic levels. It has helped in deciding the systematic
position of primitive vesselless families such as Amborellaceae, Tetracentraceae, Trochodendraceae
and Winteraceae, all of Magnoliales of Angiosperms. Phylogenists have suggested that because of
the presence of specialised wood, Amentiferae cannot be considered primitive. Amentiferae includes
families, such as Fagaceae, Juglandaceae and Betulaceae.
102 Plant Taxonomy
Wood anatomy has also provided evidence for assigning a definite position to taxa of uncertain
affinity. Placement of Myristicta close to Lauraceae, delimitation of different subgenera of Quercus
of Fagaceae, and non-inclusion of Calycanthaceae in Rosales or Myrtales are all supported by the
wood anatomy.
The type and arrangement of vascular bundles, type of secondary growth, and different characters
of xylem and phloem elements have been used in solving phylogenetic problems and delineating
different taxa.
Some important wood elements of taxonomic importance are undermentioned:
1. Vessel elements with pittings on their lateral walls, vessel abundance, presence of solitary
or aggregate vessel groupings, sculpturing on vessel walls, and distribution of vessels are
characters of systematic value.
2. Vascular rays, their width, dimensions, abundance and degree of their wall thickness are
useful taxonomic criteria.
3. Axial parenchyma, its distribution, arrangement, length and width of its cells, thickness and
lignification of its wall, and its absence are all characters of taxonomic significance.
4. Presence or absence of stroried wood is an important taxonomic character.
5. Presence or absence of latex vessels, resins, gums, crystals, etc. in the wood are also the
characters of taxonomic importance.
9.3 FLORAL ANATOMY IN RELATION TO TAXONOMY

The importance of floral anatomy in relation to taxonomy has been described by several workers,
including Eames (1953), Puri (1952, 1958, 1962), and Murty and Puri (1980). In majority of these
studies, main emphasis is laid on the path and distribution of vascular bundles within the receptacle
and floral parts.
Puri (1954) has shown strong affinities between Passifloraceae and Moringaceae on the basis of
the orientation of placental strands with reference to floral axis. On the basis of floral anatomy,
Kale and Pai (1979) have supported the removal of genus Trichopus from family Dioscoreaceae and
creation of a new family Trichopodiaceae, with Trichopus as its sole representative.
Floral anatomical studies have been useful in solving some fundamental questions like the nature
of flower, carpel, inferior ovary, and also several problems related with homologies, phylogeny and
taxonomy. Using clearing techniques, Melville (1960) investigated the floral vasculature of large
number of angiosperms and proposed the gonophyll theory of the origin of angiosperm flowers.
According to Puri (1958) the structure of the gynoecium is essentially alike in Capparidaceae,
Cruciferae, Moringaceae and Papaveraceae. The placentation in these families is parietal, but the
placental strands are inverted and occur on the inner side of the secondary marginal bundles pos-
sessing normal orientation. This study has given the support to the view that parietal placentation has
been derived from axile placentation, and Moringaceae should be included in order Rhoeadales. On
the basis of floral anatomical studies, Roy (1949) proposed the removal of Trapa from Onagraceae
and its inclusion in a separate family Trapaceae. Eyde (1975) interpreted the vestigial organs and
modified parts of flowers by studying the floral anatomy of several taxa.
Contribution of floral anatomy in resolving the taxonomic position of some disputed taxa is dis-
cussed below:
1. Floral anatomy of Annonaceae, Calycanthaceae and Menispermaceae confirms that all these
families originated from Ranunculaceae.
2. Floral anatomy of Polemoniaceae and Caryophyllaceae suggest that the former have been
derived from a caryophyllaceous stock.
3. Uniformity in floral vasculature of Solanaceae and Scrophulariaceae suggests that both should
be included in one single order, Scrophulariales.
4. Formerly, Cyperaceae and Gramineae were treated together in one single order. But
Hutchinson (1973) treated them separately in Cyperales and Graminales. Floral anatomical
studies of both the families support their separation as suggested by Hutchinson.
5. Separation of Paeonia from Ranunculaceae and its inclusion under a separate family
Paeoniaceae was supported by the floral anatomical studies.
6. Transfer of Hydrocotyle asiatica L. to the genus Centella in the form of Centella asiatica
L. was confirmed by studies of floral anatomy.
7. Systematists have suggested that there is no close affinity between Cyrtandromoea and
other members of Gesneriaceae, and it should be transferred to Scrophulariaceae. Floral
anatomical studies also show a close relationship between Cyrtandromoea and members
of Scrophulariaceae because of the presence of several lateral traces in carpels, a bilocular
ovary, and absence of a disc in both.
8. Lilaea, earlier included under family Scheuchzeriaceae, was later separated in an indepen-
dent family Lilaeaceae by systematists. Floral anatomy supported the removal of Lilaea
from Scheuchzeriaceae because both differ in their vascular supply of flower and number of
ovules.
9.4 CYTOLOGY IN RELATION TO TAXONOMY, OR CYTOTAXONOMY

Study of cell is referred as cytology. But according to Solbrig (1968) and Stebbins (1971), only the
details of the chromosomes have been used in resolving many taxonomic problems. Utilization of
the characters and phenomena of cytology for the explanation of taxonomic problems is referred as
cytotaxonomy.
The term karyotype is used for the phenotypic appearance of the somatic chromosomes. The
diagrammatic representation of the karyotype is termed as idiogram.
The characteristics of the chromosomes, which have proved to be of taxonomic value, include
(1) chromosome number, (2) chromosome size, (3) chromosome morphology, and (4) chromosome
behaviour during meiosis.
9.4.1 Chromosome Number

Usually, all individuals within a species possess the same chromosome number. In angiosperms,
the haploid chromosome number varies between n = 2 (Haplopappus gracilis of Compositae) and
n = 132 (Poa littorosa of Gramineae). However, the majority of them show a range between n = 7
104 Plant Taxonomy
and n = 12. The highest chromosome number recorded for vascular plants is found in a pteridophyte,
Ophioglossum reticulatum (2n = 1260).
Variation or constancy in the chromosome number, within taxa of different categories, prove to
be important characters for taxonomic groupings.
The chromosome number in some plants remains constant in all species, e.g. all species of Pinus
and Quercus possess n = 12 chromosomes. Such numbers are called constant chromosome numbers.
The species bearing constant chromosome numbers are called homoploids. These numbers are, of
course, of limited importance, but prove useful in knowing a particular genus. In several genera of
vascular plants polyploid series are present. Polyploids are the plants which possess higher chromo-
some numbers because of the multiplication of genomes or chromosome sets, e.g. different species
of Aster have n = 9 or n = 18 or n = 27, etc. Such a series of polyploidy, in which the chromosome
numbers of a taxon are in the proportion of its exact multiples, is called euploidy. On the other hand,
if the chromosome numbers of a group bear no simple numerical relationships with each other, the
series is called aneuploidy, e.g. different species of Brassica bear n = 6, 7, 8, 9, or 10.
9.4.2 Chromosome Size

The chromosome size has been very useful in understanding relationships in several taxa. In most
plants, the length of a chromosome varies from 0.5 to 30 µ. Among monocots, the members of
Zingiberaceae possess small chromosomes, of Iridaceae small to medium-sized, of Amaryllidaceae
large-sized, while those of Liliaceae possess chromosomes of varying sizes. Except Eleocharis and
Fimbristylis, all the members of Cyperaceae possess extremely small chromosomes. According to
Stebbins (1938) the chromosome size is characteristic of only certain groups and families, and not
related to the phylogeny of angiosperms.
9.4.3 Chromosome Morphology

Usually the chromosomes are characterized as median, submedian, subterminal or terminal with
reference to their length and the position of the centromere. The centromere location marks the
position of the primary constriction. Additional constrictions are called secondary constrictions.
Occasionally, a secondary constriction may be present near the terminal end of a chromosome,
separating its small segment called satellite. The chromosomes may be symmetrical or asymmetrical.
Symmetrical ones possess two equal arms and a median centromere. Asymmetrical ones possess
unequal arms and subterminal centromeres.
Relative length of the arms of the chromosomes, position of the centromere, presence of satellites,
etc. are some characters of taxonomic significance.
9.4.4 Chromosome Behaviour at Meiosis

Significant taxonomic information may be obtained by the study of the behaviour of the chromo-
somes during meiosis. Degree of sterility and occurrence of hybridisation are determined by the
behaviour of chromosomes during meiosis. Abnormalities in meiosis, such as non-pairing, crossing
over, unequal interchanges or translocations, bridge formation, lagging chromosomes, etc. have all
proved to be of systematic value.
9.4.5 Systematic Value of Cytological Studies

According to Jackson (1971) chromosome characteristics have been found to be useful at all taxo-
nomic levels. A few of such common characteristics of systematic value are undermentioned:
1. Members of Cyperaceae and Juncaceae possess chromosomes with diffuse or non-localized
centromeres, and also show inverted meiosis. This reflects a close association between these
two families.
2. In Ranunculaceae, reshuffling of the genera has been done in the light of cytological data.
All the genera with n = 7, 8 and 9, as well as genera with long chromosomes and short
chromosomes are accommodated in tribes Anemoneae and Helleboreae. In Thalictrum and
Anemonella of Anemoneae, and Isopyrum and Aquilegia of Helleboreae, the base number is
n = 7 and chromosomes are small-sized. Because of such a karyological similarity, all these
genera have now been grouped in tribe Thalictreae. In the same way, genera Coptis and
Zanthorhiza of Helleboreae have n = 9 and very small chromosomes and therefore grouped
under Coptideae.
3. Yucca had long been treated as a member of Liliaceae because of the superior ovary, and
Agave of Amaryllidaceae because of the inferior ovary. Hutchinson shifted both Yucca
and Agave to Agavaceae. The presence of bimodal karyotype with 25 small and 5 large
chromosomes in both of them, justified their placement in one single family, as done by
Hutchinson.
4. In the subfamily Bambusoideae of Gramineae n = 12, and in the subfamily Poideae n = 7.
This indicates that the chromosome numbers have proved to be of taxonomic utility also at
the subfamily level. Stebbins (1958) provided information on the evolution of grasses on the
basis of cytogenetics.
5. The hexaploid wheat (Triticum aestivum) has a genome constitution of AABBDD. In this
case the genome A has been contributed by Triticum monococcum, genome B by Aegilops
speltoides, and genome D by Aegilops squarrosa. This shows that by such a study of meiosis
the parentage of many polyploid taxa may be located.
6. The basic chromosome number in Loranthaceae is n = 9 while in Viscaceae there is a series
of aneuploid numbers ranging between 10 and 14. Wiens (1975), therefore, suggested their
separation from each other on the basis of the cytological evidence.
7. Delimitation of the tribes in Compositae has been done on the basis of chromosome
numbers.
8. On the basis of cytological studies, Lewis (1951, 1953) submerged the genus Godetia in
Clarkia (Onagraceae).
9. Naik (1977) differentiated three species of Chlorophytum of Liliaceae on the basis of cyto-
logical data. According to him C. bharuchae has 2n = 16 chromosomes while C. glaucum
and C. glaucoides have 2n = 42. Both the latter species, having 2n = 42, differ in their
karyomorphology.
10. Warburg (1938) studied taxonomy of Geraniales on the basis of cytological studies.
11. Manton (1932) confirmed the formation of subdivisions of Brassicaceae on the basis of cyto-
logical studies. All the subfamilies have different base chromosome numbers.
106 Plant Taxonomy
12. Genus Cistus (Cistaceae), formerly included in Helianthemum, has chromosome number 8
while Helianthemum has base chromosome number 9. So, Cistus should be recognized as a
separate genus.
13. Genera Physaria and Lesquerella of Brassicaceae were treated as single genus by several
taxonomists. Their different cytological details suggest them to be treated as two different
genera.
14. A new classification of the genus Narcissus of Amaryllidaceae has been proposed by
Fernandes (1951) on the basis of cytological studies.
15. Stebbins (1971) distinguished grasses on the basis of size and number of chromosomes.
According to him bambusoid grasses have small and many chromosomes, chloridoid grasses
have small and few chromosomes, festucoid grasses have large chromosomes, and panicoid
grasses have medium-sized chromosomes.
16. According to Mahabale and Cheenaveeraiah (1953) the palm species with n = 16 chromo-
somes have pinnate leaves while the palm species with n = 18 have palmate leaves. However,
there exist exceptions in both the cases.
17. Cheenaveeraiah (1962) suggested on the basis of karyotypic studies of species of Aegilops
that its section Sitopsis should either be shifted from Aegilops to Triticum or it should be
given the rank of a new genus.
18. Sharma (1956), on the basis of his studies of Araceae, Amaryllidaceae and Dioscoreaceae,
proposed that the changes in karyotypes of somatic tissue play a distinct role in evolution. He
further proposed (Sharma, 1964) that large chromosomes, low chromosome number and sym-
metrical karyotype represent a primitive status, while small chromosomes, high chromosome
number and extreme asymmetry of karyotype represent the advance status. These principles
provided interesting results in taxonomy of Alismataceae, Liliaceae, Amaryllidaceae and
Dioscoreaceae.
9.5 PALYNOLOGY IN RELATION TO TAXONOMY

9.5.1 Pollen Characters and Related Studies
Study of pollen and spores is called palynology. The shape and symmetry of pollen grain, the archi-
tecture of its wall, exine stratification, sculpture and structure; and type, number, position, shape
and structure of its aperture, are some of the basic characters, which prove useful at all taxonomic
levels.
Palynological characters have been used in solving several taxonomic problems, including the repo-
sitioning of several disputed taxa, and interpretation of problems relating to the origin and evolution
of different groups (Nair, 1980). Cronquist (1981) and several other workers have made the exclusive
use of pollen characters in providing classification of angiosperms. Erdtman (1963) used the pollen
characters in discussing and solving the taxonomic problems of 105 families. Pollen morphology
has proved to be of great value in the classification of Compositae (Stix, 1960), Acanthaceae (Raj,
1961), Euphorbiaceae (Kohler, 1965), Gentianaceae (Nilsson, 1967), Saxifragaceae (Nair, 1970),
Asclepiadaceae (Radford, et al., 1974), Caryophyllidae (Cronquist, 1981), etc. Heywood (1967) has
gone up to the extent of stating that exine details of pollen are such that they can be used in plant
identifications much in the way that fingerprints are used for the identification of criminals.
9.5.2 Monosulcate and Tricolpate Pollen

Jones and Luchsinger (1987) mentioned that angiosperms contain two basic kinds of pollen grains:
monosulcate and tricolpate. Monosulcate pollen grains are characteristic of primitive dicotyledons,
several monocotyledons, pteridosperms, and cycads. Such pollen grains are boat-shaped in outline
and possess one long germinal furrow and a germinal aperture. Tricolpate pollen grains are charac-
teristic of advanced dicotyledons. Such pollen grains have three germinal apertures and are globose
in shape.
9.5.3 Stenopalynous and Eurypalynous

Such a taxon, in which the type of the pollen is constant and characteristic, is called stenopalynous
or unipalynous, e.g. Gramineae is unipalynous, having monoporate aperture and somewhat smooth
exine surface. On the other hand, such taxa, in which pollen types vary in size, aperture, exine
stratification, etc., are called eurypalynous or multipalynous, e.g. Euphorbiaceae is multipalynous
with grains of various aperture forms.
Cruciferae, Labiatae and Asclepiadaceae are unipalynous families while Acanthaceae, Rubiaceae,
Compositae and Verbenaceae are highly multipalynous families.
9.5.4 NPC-System
The classification of pollen is based on the number-position-character analysis, called NPC-system.
Palynological studies suggest that the taxa with the same general NPC-formula be grouped together,
and those showing different NPC, separately. NPC-system helps in providing a three-dimensional
classification, and also in the preparation of diagnostic keys below the family level. Cronquist (1981)
also emphasized on the importance of the NPC-system in solving several taxonomic problems. For
example, the pollen grains of Parietales (as investigated by means of “palynological compass needle”)
are 3-tremes (N3), zonotreme (P4) and colporate (C5). Therefore, the NPC-formula for Parietales is
345.
9.5.5 Some Other Pollen Characters of Systematic Value

1. Size and shape of pollen are important diagnostic characters. The shape may be circular,
triangular, etc. in polar view, whereas it may be prolate, perprolate, spheroidal, subpolar, etc.
in equatorial view.
2. Apertures may be proximal (gymnosperms and monocotyledons), distal (pteridophytes), zonal
(dicotyledons) or global (dicotyledons).
3. Exine sculpturing may be of depression type (e.g. reticulate, foveolate, striate, fossulate, areo-
late, scrobiculate, etc.) or excrescence type (e.g. granulose, verrucate, spinulose, tuberculate,
spinose, gemmate, clavate, etc.).
4. Pollen associations, such as their arrangement in tetrads, presence of pollinia, etc. are char-
acteristic of several angiospermic families.
108 Plant Taxonomy
5. Number of nuclei in the pollen grains at the time of their release is a valuable taxonomic
and phylogenetic character.
9.5.6 Some Examples of Role of Palynology in Taxonomy

1. Author (Sharma, 1987) investigated the pollen morphology of 16 Indian species of Cyperus,
and prepared a key to differentiate all of them on the basis of pollen characters. Of the 16
investigated species, the pollen grains of six (Cyperus iria, C. difformis, C. squarosus, C.
triceps, C. flabelliformis and C. paniceus var. roxburghianus) are predominantly l-colpate,
two (C. exaltatus and C. pumilus) are predominatly 2-colpate, four (C. rotundus, C. laevi-
gatus, C. alulatus and C. bulbosus) are predominantly 4 aperturate with 3 colpi and 1 pore,
three (C. compactus, C. kyllingia and C. globosus) are predominantly pantoaperturate, and
one (C. digitatus) is predominantly 1-porate. Individual species are also differentiated on the
basis of pollen characters.
2. On the basis of apertural morphoforms of pollen, Nair (1974) proposed a palynological
classification. He divided the entire plant kingdom into (i) Amorphosporophyta, including
thallophyta with no apertural markings, (ii) Trimorphosporophyta, including Archegoniatae
with monolete, trilete, or alete forms, and (iii) Polymorphosporophyta, including angiosperms
with different forms of apertural types.
3. According to Meyer (1975) gymnosperms contain alveolar or granular ektexine and laminated
endexine, whereas angiosperms contain columnar or granular ektexine and nonlaminated
endexine.
4. A massive exine and thin intine is present in angiosperm pollens. But in certain taxa
among monocots (e.g. Musaceae, Amaryllidaceae, Cannaceae, Zingiberaceae, Costaceae and
Heliconiaceae), the exine is highly reduced and the intine is well-developed.
5. Several angiospermic taxa have distinctive pollen types. Gramineae have smooth sulcate
pollen, Malvaceae and Compositae have typically spinulose exine, and Plumbaginaceae have
verrucate exine. Pollens are binucleate in Magnoliidae and trinucleate in Caryophyllidae.
6. The exine pattern is useful in recognising different species of a genus. According to Nair
(1974) pollens are pilate in Bauhinia acuminata, striate in B. krugii, verrucate in B. retusa,
reticulate in B. racemosa, spinulate in B. malabarica, and reticulate tuberculate in B.
purpurea.
7. Phylogenetic relationships are determined using pollen characters in several cases. Genus
Cicer is usually placed under the tribe Viceae of Papilionaceae. Palynological studies of Cicer
suggest that it is more close to the tribe Ononideae and should be transferred from Viceae
to Ononideae (Clarke and Kupicha, 1976).
8. Pollen grains are associated in tetrads in several families of dicots (Annonaceae, Winteraceae,
Mimosaceae, Droseraceae, etc.) and monocots (Cyperaceae and Juncaceae). In Asclepiadaceae
and Orchidaceae, they form pollinia.
9. Elastic threads join massulae in the pollinia in Orchidaceae (Vijayraghavan and Shukla,
1980). Viscin threads of similar nature are found in Onagraceae.
10. According to Kooiman (1972) 2-nucleate pollen grains of Lamiaceae are tricolpate while
3-nucleate pollen grains are 6-colpate.
11. Palynological evidences have supported the separation of Paeoniaceae from Ranunculaceae,
Nelumbonaceae from Nymphaeaceae, Fumariaceae from Papaveraceae, and Bombaceae from
Malvaceae.
12. Removal of Podophyllum from Berberidaceae and its inclusion in a separate family
Podophyllaceae is effected on palynological grounds.
13. Pollen characters prove useful in distinguishing the genera Salix and Populus of Salicaceae,
Phytolacca and Rivinia of Phytolaccaceae, and several genera of Betulaceae, Primulaceae
and Acanthaceae.
14. According to Nair (1980), three well-defined evolutionary stocks among angiosperms (i.e.
Monocot stock, Magnolian-dicot stock, and Ranalian-dicot stock) form the palynological basis
of the triphyletic theory of angiosperms.
15. The genus Krameria, variously placed under Leguminosae, Polygalales, etc. was recom-
mended to be placed in a separate monotypic family Krameriaceae by Simpson and Skvarla
(1981) on palynological grounds.
16. Palynological studies suggest that Thunbergioideae of Acanthaceae should be given a family
status.
9.6 EMBRYOLOGY IN RELATION TO TAXONOMY

Embryology is the study of micro- and megasporogenesis, gametophyte development, fertilization,
and development of endosperm, embryo, and seed coats. Embryological evidences have been used
in solving the taxonomical problems at almost all levels. These evidences have resolved the doubtful
systematic positions of several taxa. However, the role of embryology in solving taxonomic problems
was first brought into prominence by a German embryologist, Schnarf in 1931. And in the later
years, excellent detailed treatments of the role of embryology were published by Johansen (1950),
Maheshwari (1950, 1964), Kapil (1962), Johri (1963, 1967), Davis (1966), Sporne (1969), Philipson
(1974), Dickison (1974), Pa1ser (1975), Bhojwani and Bhatnagar (1979), Kapil and Bhatnagar (1980),
etc. According to Jones and Luchsinger (1987), the embryological characters have proved to be of
significant help “in determining relationships within families, genera and species”, and have proved
to be less useful “at the rank of order, subclass, or class”.
9.6.1 Basic Embryological Evidentiary Characters

According to Maheshwari (1950, 1964), Bhojwani and Bhatnagar (1979) and Radford (1986), some
basic embryological characters which have proved to be of special importance in taxonomic con-
siderations include (i) presence and type of anther tapetum; (ii) number and arrangement of anther
loculi; (iii) type of anther endothecium; (iv) quadripartition of microspore mother cell; (v) mature
pollen grains; (vi) development, structure, position, vasculation, and orientation of ovule; (vii) origin
of sporogenous tissue in ovule; (viii) megasporogenesis and development of embryo sac; (ix) presence
of aril; (x) form of embryo sac; (xi) fertilization; (xii) type of embryo; (xiii) type of embryogeny; (xiv)
endosperm formation; (xv) type of haustorium formation; (xvi) seed-coat; (xvii) cotyledons, etc.
110 Plant Taxonomy
9.6.2 Some Examples of Role of Embryology in Taxonomy

A lot of literature has been published on the role of embryology in solving taxonomic problems dur-
ing the last 40 years. It is not possible to review it entirely. Some examples are undermentioned:
1. Dicots and Monocots Angiosperms are universally divided into dicotyledons and mono-
cotyledons. This primary classification of angiosperms is based on one major embryological
character, i.e. number of cotyledons.
2. Caryophyllales Embryological characters, such as trinucleate pollen, bitegmic crassinu-
cellate ovules which are campylotropous or amphitropous, seed with peripheral embryo,
and perisperm with little or no endosperm, are the characters which are found only in
Caryophyllales, more widely known as Centrospermae (Cronquist, 1968).
3. Helobiae This monocotyledonous order, treated as a subclass in some recent systems of
classification, is characterised by the presence of helobial type of endosperm.
4. Orchidales The distinguishing embryological character of the members of this order is the
presence of undifferentiated embryo and very little or no endosperm.
5. Podostemaceae Members of this family are recognised because of the formation of pseu-
doembryo sac which is formed by the disintegration of nucellar cells below the embryo sac.
The presence of paired pollen grains, tenuinucellate ovules and prominent suspensor haus-
toria, and absence of antipodals and triple fusion are other characteristics of the family.
6. Onagraceae The family is recognized by the presence of Onagrad-type of embryo-sac.
Such an embryo sac is derived from the micropylar megaspore of the tetrad. The lower three
chalazal megaspores do not disintegrate. In this type of embryo sac, antipodals are absent.
7. Cyperaceae In flowering plants, four functional microspores develop from each microspore
mother cell. But in Cyperaceae, each microspore mother cell gives rise to only one pollen
grain. Out of the four nuclei, formed by the meiosis of a microspore mother cell, three are
cut off on one side and do not form pollen grains. The remaining fourth nucleus divides
mitotically to give rise to a vegetative cell and a generative cell. Mitosis in the generative
cell results in the formation of two sperm cells.
8. Lemnaceae Phylogenetic studies indicate that Lemnaceae have been derived either from the
Helobiales or from the Araceae. But on the basis of the embryological studies, Maheshwari
(1954, 1958) suggested that Lemnaceae have been evolved from Araceae and not from
Helobiales.
9. Crassulaceae Embryological studies of Crassulaceae suggest that it should be placed in the
order Rosales close to the family Saxifragaceae.
10. Loranthoideae and Viscoideae Members of the subfamily Loranthoideae, with triradiate
pollen, Polygonum-type embryo sac and composite endosperm show the presence of suspensor
and polyembryony. On the other hand, members of the subfamily Viscoideae contain spheri-
cal pollen, Allium-type embryo sac, non-composite endosperm and show the general absence
of suspensor and polyembryony. These differences clearly suggest that both the subfamilies
should be elevated to the rank of families, i.e. Loranthaceae and Viscaceae.
11. Paeonia It has been treated by most of the taxonomists as a member of the monogeneric
tribe Paeonieae of the family Ranunculaceae. But Paeonia differs from Ranunculaceae in its
chromosome number, vascular anatomy, floral anatomy as well as in embryological details.
It shows a unique embryogeny. Its young zygote first becomes coenocytic and then the cell
formation starts from its periphery. From some of the peripheral cells develop the embryo
initials. Paeonia bears arillate seeds and a follicle fruit while in Ranunculaceae the seeds
are non-arillate and the fruit is an achene. All these details suggest that Paeonia belongs to
an independent family Paeoniaceae.
12. Exocarpus Because of the presence of a naked ovule and pollen chamber, Exocarpus
was removed from Santalaceae of angiosperms and was treated as a member of the family
Exocarpaceae near Taxaceae in gymnosperms. But the presence of a typical angiospermic
flower, Polygonum-type of embryo sac, cellular endosperm and some other embryological
characters, led Ram (1956) to confirm that Exocarpus belongs to the family Santalaceae of
angiosperms, and not with gymnosperms.
13. Trapa Majority of the taxonomists treat Trapa as a genus of Onagraceae while others con-
sider it to belong to the family Hydrocaryaceae. But its embryological details (Polygonum-
type embryo sac, absence of endosperm, well-developed suspensor haustorium, extremely
reduced one cotyledon, etc.) suggest that Trapa should be treated under an independent family
Trapaceae.
14. Pentaphragma It has been treated under the families Campanulaceae, Boraginaceae and
Pentaphragmataceae by different workers. But its similarities with Campanulaceae in its
embryological details (i.e. ovule, endothecium, embryo sac, endosperm and embryogeny)
deserve its final inclusion in this family.
15. Butomus The presence of the Polygonum-type of embryo sac in Butomus and the Allium-
type of embryo sac in the other genera of the family Butomaceae, suggest that only Butomus
should be retained in Butomaceae. The other genera of this family should be transferred
either to Alismataceae or to Limnocharitaceae.
16. Parnassia It is generally treated as a member of the family Saxifragaceae. But the total
dissimilarity of its embryological details with the other genera of Saxifragaceae suggests its
removal from Saxifragaceae and inclusion in a separate family Parnassiaceae.
17. Peganum It has been variously treated as a member of Rutaceae or Zygophyllaceae. But
its embryological details resemble those of Linaceae. The recent embryological findings,
therefore, suggest its inclusion in an independent family, Peganaceae near Linaceae.
All the abovementioned examples confirm that the embryology plays a definite and significant
role in solving taxonomic problems.
9.7 CHEMISTRY IN RELATION TO TAXONOMY 1

Application of chemistry to taxonomy is called chemical taxonomy or Chemotaxonomy 1. Chemical
evidences are used in determining the relationship among taxa of different categories. Some of the
1
For details of chemotaxonomy, see Chapter 11.
112 Plant Taxonomy
major classes of the chemical evidence include flavonoids, alkaloids, amino acids, fatty acids, aro-
matic compounds, terpenoids, polysaccharides, carotenoids, etc.
Cronquist (1981) cited following examples to indicate the use of chemistry in solving taxonomic
problems:
1. Caryophyllales produce betalains and not anthocyanins.
2. Polygonales produce anthocyanins and not betalains.
3. Juglandales are aromatic plants while Fagales are non-aromatic.
4. Highly aromatic compounds are found in Lamiaceae.
5. Alkaloids are very common in Solanaceae.
6. Sapindaceae have plenty of tannins.
9.8 ECOLOGY IN RELATION TO TAXONOMY

Ecological evidences also contribute to our understanding of several taxonomic problems. Some of
the basic ecological evidentiary characters include habitat (water, sand, mud, rock, marsh, desert,
etc.), abiotic relations (light, moisture, climate, soil, etc.), biotic relations (symbiotic, nutritional,
reproductive, etc.), spatial relations (areal, distributional, etc.), and temporal relations (succession,
phenology, periodicity, etc.)
Ecological evidences prove helpful in understanding the distribution of taxa, variation within taxa,
and adaptations of plants. These evidences have also been used for providing an explanation about
the origin of the flowering plants, and also about the development of classification of several taxa,
mainly below the level of the genus. Ecological studies also show that many morphological features
of plants have a direct correlation with the environmental factors, such as light, humidity, water, etc.
Ecologists make a substantial contribution to the systematics by examining ecotype variation, soil
specializations, seedling establishment ecology, seed dispersal mechanism, etc.
9.9 PALAEOBOTANY IN RELATION TO TAXONOMY

Information regarding the correct evolutionary history of the present-day plants is gathered from
the fossil records or palaeobotany. Any theory of evolution of plants is incomplete without any
palaeobotanic evidence. Both, microfossils (e.g. pollen) and macrofossils (e.g. stems, leaves, etc.)
are used as the source of the systematic data. Palaeobotanic studies prove helpful in determining
the evolution of the floras of the past, in the comparative morphology, as well as in determining
the ecological conditions of the past.
According to Beck (1976) and Taylor (1981) the palaeobotanic studies are helpful in the phylog-
eny of angiosperms. Dilcher (1979) is of a firm opinion that the palaeobotany provides important
information about several aspects of the early history of angiosperms, as well as their origin and
diversification. Number of new concepts have been developed about the primitive flower types by
Dilcher (1979) on the basis of his palaeobotanic studies. These studies also suggest that diversified
representatives of the modern Amentiferae were commonly present in Middle Eocene. Palaeobotanists
are also now of a firm opinion that the Middle Eocene flora may provide definite information about
the origin of plants in general and the evolution of flower in particular.
9.10 ELECTRON MICROSCOPY IN RELATION TO TAXONOMY

Electron microscopy has brought revolution in all biological fields, and so it has its implications
on taxonomy too. Ultrastructural studies, which include scanning electron microscopy (SEM) and
transmission electron microscopy (TEM), have provided definite clues to the phylogeny of angio-
sperms. Pollen grains, trichomes, spores, seeds and plant surface have been investigated in detail
using electron microscopy.
The taxonomic value of sieve-element plastids in the classification of flowering plants was dem-
onstrated by Behnke (1972) using transmission electron microscopy. Two classes of plastids were
recognized, i.e. s-type and p-type. The s-type plastids accumulate starch while p-type accumulate
only proteins or starch and protein both. Valuable taxonomic information is provided by TEM studies
in certain taxa. Different types of wall structure in some members of Compositae, dilated cisternae
of endoplasmic reticulum in some genera of Cruciferae and Capparidaceae, and presence of crystal-
loids in the cotyledon mesophyll cells in Cucurbitaceae are some characters of systematic value.

1. Explain in detail the role of external morphology in relation to taxonomy.
2. Give a brief account of some modern trends in plant taxonomy.
3. In determining relationship between different genera, families, orders and other taxa, ana-
tomical characters are most useful. Explain with examples.
4. Describe the importance of floral anatomy and cytology in relation to taxonomy.
5. Explain in brief the role of embryology in relation to taxonomy.
Suggested Reading
Bailey, I.W., 1957, Wood anatomy in the study of phylogeny and classification angiosperms, J. Arnold
Arbor. Harv. Univ. 38: 243–54.
Behnke, H. D., 1972, Sieve tube plastids in relation to angiosperm systematics: An attempt towards a clas-
sification by ultrastructural analysis, Bot. Rev. 38: 155–97.
________ 1977, Transmission electron microscopy and systematics of flowering plants, Pl. Syst. Evol. Suppl.
1: 155–78.
Bendz, G. and J. Santesson, 1973, Chemistry in botanical classification, Proc. 25th Nobel Symp. Acad.,
New York.
Cole, G. T. and H.D. Behnke, 1975, Electron microscopy and plant systematics, Taxon 24: 3–15.
Davis, G.L., 1966, Systematic Embryology of the Angiosperms, Wiley, New York.
Davis, P.H. and V.H. Heywood, 1965, Principles of Angiosperm Taxonomy, Van Nostrand, Princeton,
N.J.
Dickison, W.C., 1975, The bases of angiosperm phylogeny: Vegetative anatomy, Ann. Mo. Bot. Gard. 62:
59–62.
114 Plant Taxonomy
Erdtman, G., 1966, Pollen Morphology and Plant Taxonomy, Vol. I. Angiosperms, Hafner, New York.
Eyde, R.H., 1975, The bases of angiospenn phylogeny: Floral anatomy, Ann. Mo. Bot. Gard. 62: 521–37.
Gibbs, R.D., 1974, Chemotaxonomy of Flowering Plants, 4 Vols. McGill-Queens Univ. Press, Montreal.
Harborne, J.B. and B.L. Turner, 1984, Plant Chemosystematics, Academic Press, New York.
Howard, R.A., 1963, The vascular structure of the petiole as a taxonomic character, Adv. Hort. Sci. 3:
7–13.
Jackson, R.C., 1971, The karyotype in systematics, Ecol. Syst. 2: 327–68.
Johri, B.M., 1963, Embryology and taxonomy (In) Recent Advances in the Embryology of Angiosperms,
Maheshwari, P. (ed.). International Soc. Pl. Morphology, New Delhi.
________ 1967, Angiosperm’s embryology and taxonomy, Bull. Natn. Inst. Sci. India 34: 263–68.
Jones, S.B. Jr. and A.E. Luchsinger, 1987, Plant Systematics (2nd ed.), McGraw-Hill, Singapore.
Kapil, R.N., 1962, Some recent examples of the value of embryology in relation to taxonomy, Bull. Bot.
Surv. Ind. 4: 57–66.
Kruckeberg, A.R., 1969, The implications of ecology for plant systematics, Taxon 18: 92–120.
Maheshwari, P., 1964, Embryology in relation to taxonomy, (In) W.B. Turrill (ed.) Vistas in Botany IV.
Macmillan, New York.
Metcalfe, C.R., 1964, An anatomist’s view on angiosperm classification, Kew Bull. 9: 427–40.
________ and L. Chalk, 1950, Anatomy of the Dicotyledons, Vols. I. & II, Oxford Univ. Press, Oxford.
Murty, Y.S. and V. Puri, 1980, Floral anatomy in relation to taxonomy, Glimp. Pl. Res. 5: 222–243.
Nair, P.K.K., 1980, Glimpses in Plant Research V. Modern Trends in Plant Taxonomy, Vikas Publ., New
Delhi.
Paliwal, G.S. and S.K. Anand, 1978, Anatomy in relation to taxonomy, Acta Bot. Indica 6: 1–20.
Philipson, W.R., 1974, Ovular morphology and the major classification of dicotyledons, Bot. J. Linn. Soc.
68: 89–108.
Puri, V., 1952, Floral anatomy in relation to taxonomy, Agra Univ. J. Res. (Sci) 1: 15–35.
________ 1958, Floral anatomy and taxonomy, Ind. Bot. Ser. Mem. 1: 15–18.
________ 1962, F1oral anatomy in relation to taxonomy, Bull. Bot. Surv. India. 4: 161–65.
Sharma, A.K., 1954, Cytology as an aid in taxonomy, Bull. Bot. Soc. Bengal 18: 1–4.
Sharma, O.P., 1987, Pollen morphology of Cyperus, Bangladesh J. Bot. 16(2): 141–50.
________ and R. Shiam, 1981a, Occurrence of cuticular papillae in Cyperus, Curr. Sci. 50(5): 236.
________ and R. Shiam, 1981b, Pore formation in Cyperaceae: A new report, Curr. Sci. 50(2): 452–53.
________ 1984, Epidermal structures of culm in Cyperus with a discussion of silica bodies in Cyperaceae,
Bangladesh J. Bot. 13(1): 16–24.
Smith, P.M., 1976, The Chemotaxonomy of Plants, Edward Arnold, London.
Stace, C.A., 1965, Cuticular studies as an aid to plant taxonomy, Bull. Br. Mus. (Nat. Hist.) Bot. 4: 1–78.
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NUMERICAL
TAXONOMY 10
10.1 WHAT IS NUMERICAL TAXONOMY?
The analysis of various types of taxonomic data by mathematical or computerized methods is called
numerical taxonomy or taximetrics. This approach of systematics involves the numerical evaluation
of the similarities or affinities between taxonomic units, and then arrangement of these units into
taxa on the basis of their affinities.
Adanson (1763), a French botanist, was the first to put forward a plan for assigning numerical
values to the similarity between organisms. He tried to use as many characters as possible for the
classification, and such classifications were recognised as Adansonian classifications. Adansonian
principles have developed several new methods in taxonomy during recent years, particularly after
1960, and these methods are all included under numerical taxonomy.
The use of modern electronic data-processing techniques and several other methods of the numeri-
cal taxonomy have helped in the evolution of several new classifications of plants during the past
40 years. Several objectives and qualitative methods are employed in their evolution.
According to Heywood (1967) the numerical taxonomy may be defined as “the numerical evalu-
ation of the similarity between groups of organisms and the ordering of these groups into higher
ranking taxa on the basis of these similarities.”
10.2 PRINCIPLES OF NUMERICAL TAXONOMY

Numerical taxonomy is based on certain principles which are also called neo-Adansonian principles.
Sneath and Sokal (1973) have enumerated following seven principles of numerical taxonomy:
1. The greater the content of information in the taxa of a classification system and more the
characters on which it is based, the better a given classification will be.
2. Every character has equal weightage in creating new taxa.
3. Between any two entities, the overall similarity “is a function of their individual similarities
in each of the many characters in which they are being compared”.
116 Plant Taxonomy
4. “Distinct taxa can be recognized because correlations of characters differ in the groups of
organisms under study”.
5. Assuming some evolutionary mechanisms and pathways, the phylogenetic conclusions may
be drawn from the taxonomic structure of a group and from character correlations.
6. The science of taxonomy is viewed and practiced as an empirical science.
7. Phenetic similarity is the base of classifications.
10.3 LOGICAL STEPS OF NUMERICAL TAXONOMY

Following are the successive steps used in the construction of taxonomic groups:
10.3.1 Operational Taxonomic Units (OTUs)

The lowest ranking taxa or the basic units in any specific study are called operational taxonomic
units or OTUs. An OTU may be an individual, a species, a genus, or a taxon of other higher rank.
Therefore, the rank of OTUs may differ from study to study. Usually, the species is treated as an
OTU.
10.3.2 Unit Taxonomic Characters or Attributes

According to Sneath and Sokal (1973) a unit character is “a taxonomic character of two or more
states, which, within the study at hand, cannot be subdivided logically, except for the subdivision
brought about by the changes in the method of coding”. Usually the phenotypic characters are used
as unit characters, e.g. the presence or absence of an awn in a spikelet of a grass.
Maximum number of unit characters, and certainly not less than 50 in any case, should be used
to obtain a fairly stable and reliable classification. Too few characters impart unreliability. A new
item of information should be contributed by each unit character. Sneath and Sokal (1973) have given
a list of several such characters which are disqualified or not useful in numerical taxonomy. Such
characters are called inadmissible characters.
Regarding the proper selection of the unit characters, Sneath and Sokal (1973) suggested that (i)
they should come from all the parts of the organisms, (ii) they should belong to all the stages of
the life cycle of the organisms, (iii) all variable characters within the group should also be used;
and (iv) due attention should be given to characters related to morphology, physiology, ecology and
distribution of the organisms.
10.3.3 Types of Basic Characters or Attributes

Following are the major types:
1. Binary Characters These characters contain two contrasting states, such as the presence or
absence of some features, e.g. presence or absence of stipules.
2. Multistate Characters These may be of following two types:
(a) Qualitative multistate characters These characters contain three or more contrasting
forms, and each form is ranked on equal footing, e.g. flower colour blue, orange or
red.
Numerical Taxonomy 117
(b) Quantitative multistate characters These characters represent measures of the size on
a continuous scale such as length, breadth, height, or weight, etc., e.g. length of a fungal
spore, or amount of mucilage produced by an alga.
10.3.4 Coding of Characters

The selected unit characters are assigned a symbol or mark, e.g. 1, 2, 3 . . ., or + and –, or 1 +, 2 +,
3 +, etc. This is called coding of characters. The symbol NC is used for “no comparison”. Coding
of characters may be of following types:
1. Two-state Coding The characters in this type of coding are divided as 1 and 0, or as +
and –. The characters recorded as 1 or + are the positive characters while those recorded as
0 or – are negative. The word NC is used if an organism is not possessing a given character
present in other organisms.
2. Multistate Coding Each quantitative multistate character may be expressed by a single
numerical value, such as 1, 2, 3, 4 . . . corresponding to the range of the variation of the
character. On the other hand, the qualitative multistate characters cannot be arranged in
a reliable sequence or definite order. Therefore, the qualitative characters are converted
conveniently into new characters, and coded as some alphabetic symbols depending on the
computer programme.
10.3.5 Estimation of Resemblance

Between any two operational taxonomic units, the resemblance is estimated or measured in terms of
similarity, i.e. percentage of characters in which they agree, or in terms of dissimilarity, i.e. percent-
age of characters in which they do not agree. In numerical taxonomy, the dissimilarity is specially
useful in the construction of taxonomic models or maps because it is the taxonomic distance between
the positions of the operational taxonomic units in the phenetic appearance.
Phenetic resemblance between taxonomic groups is usually estimated by three methods, viz.
(a) coefficients of association, (b) coefficients of correlation, and (c) measurement of taxonomic
distance between OTUs.
Coefficients of association are measured by the following formula:
S= Ns
Ns + Nd
where S is the numerical index; Ns = the number of positive features shared by any two OTUs;
and Nd = the number of positive features in one OTU and number of negative features in the other
OTU.
Coefficients of correlation (r) are measured by the following formula:
n
/[(Xij − Xi)( Xik − X k)]
rjk =
*>/ (Xij − X j) 2 H >/ (Xik − Xk) 2 H4
n n
118 Plant Taxonomy
where j and k stand for two units under comparison; Xij stands for the value of the character i in
unit j; Xik stands for the value of the character i in unit k; Xj and Xk stand for the mean for all the
characters in units j and k; n stands for number of characters.
Taxonomic distance (d) is measured by the following formula:
)/ (Xij − Xik) n2 3
n
djk =
where Xij is the character state of unit j for the character i; Xik is the character state of unit k;
n
symbol / stands for the sum over n characters; the value of the taxonomic distance (d) is the
distance in a phenetic space divided by n .
10.3.6 Cluster Analysis

Clusters are the groups of OTUs. With the help of estimation of resemblances, the affinities of differ-
ent OTUs are determined. OTUs of similar affinities are grouped together in different taxa. A taxo-
nomic system is constructed on the basis of resemblances and differences between groups of OTUs
or clusters. The main features of the cluster analysis are arranged in the form of a dendrogram.
10.3.7 Phenons and Ranks

Groups of similar organisms recognised by numerical methods are called phenons. Phenons are
equivalent to various taxonomic groups. However. the term phenon is not a synonym of the term
taxon. A phenon may or may not be equivalent to the ranks of the classical taxonomy, such as spe-
cies, genus, tribe, family, etc.
Nair (1984) has stated that “the delimitation of phenons is done by drawing a horizontal line
(Fig. 10.1) across the dendrogram at a similarity value. A line at 75%, for example, creates five
75-phenons 1; 7; 3, 5, 6; 4, 9, 10; and 2, 8; while that at 80% creates six phenons”.
10.4 ADVANTAGES OF NUMERICAL TAXONOMY

Sneath and Sokal (1973) have mentioned the following main advantages of numerical taxonomy:
1. In comparison with the conventional taxonomy, the data in numerical taxonomy are col-
lected from more variety of sources, such as morphology, physiology, chemistry, amino acid
sequences or proteins, etc.
2. Through numerical taxonomy, much taxonomic work can be done by less highly skilled
workers.
3. Numerically coded data may be used easily for the creation of keys, maps, descriptions,
catalogues, etc. with the help of existing electronic data processing systems in herbaria and
other taxonomic institutions.
4. Because the numerical methods are more sensitive in delimiting taxa they provide better keys
and classification systems in comparison to the conventional taxonomic methods.
5. The quality of conventional taxonomy is improved by numerical taxonomy as more and
better-described characters are used in the latter.
Numerical Taxonomy 119
6. Numerical taxonomy has suggested several fundamental changes in the conventional prin-
ciples of taxonomy in general and classification systems in particular.
7. Number of the existing biological concepts have been reinterpreted in the light of numerical
taxonomy.
1 7 3 5 6 4 9 10 2 8
100
90
Percent similarity
80 80% phenon line

75% ” ” ”
70
65% ” ” ”
60
50 50% ” ” ”
Fig. 10.1 Dendrogram to show formation of phenons. (From Naik, V.N. 1984, Taxonomy of
Angiosperms. Used with permission of Tata McGraw-Hill, New Delhi).
10.5 APPLICATIONS OF NUMERICAL TAXONOMY

Undermentioned are a few examples of the applications of numerical taxonomy:
1. Numerical methods are used considerably to study the similarities and dissimilarities in
bacteria, and other microorganisms.
2. Numerical taxonomy is used for delimitation of several angiospermic genera, including
Solanum (Soria and Heiser, 1961), Oryza (Morishima and Oka, 1960) and Onosis (Cook,
1969).
3. Several angiospermic taxa are reclassified using numerical methods.
4. Sneath and Sokal (1973) have mentioned several examples of the application of numeri-
cal taxonomy in several angiospermic genera, including Apocynum, Crotalaria, Cucurbita,
Chenopodium, Oenothera, Salix, Zinnia, barley cultivars, maize cultivars, wheat cultivars,
etc.

1. What is numerical taxonomy? Is it different from taximetrics?
2. Explain in brief the successive steps/logical steps of numerical taxonomy.
3. What do you mean by basic characters in numerical taxonomy? What are their main types?
Explain the meaning of “coding of characters”.
4. In relation to numerical taxonomy, write down the formulae of measuring (a) coefficients of
association, (b) coeffiecient of correlation, and (c) taxonomic distance.
5. Write in brief some applications and advantages of numerical taxonomy.
120 Plant Taxonomy
Suggested Reading
Cole, A.J., 1969, Numerical Taxonomy, Academic Press, London.
Cullen, J., 1968, Botanical problems of numerical taxonomy, (In) V.H. Heywood (ed.) Modern Methods in
Plant Taxonomy, Academic Press, London, pp. 175–184.
Rohlf, F.J. and R.R. Sokal, 1965, Coefficients of correlations and distance in numerical taxonomy, Univ.
Kansas Sci. Bull. 45: 3–27.
Sneath, P.H.A. and R.R. Sokal, 1962, Numerical taxonomy, Nature. 193: 855–860
________ and ________ 1973, Numerical Taxonomy, W.H. Freeman and Company, San Francisco.
Sokal, R.R. and P.H.A. Sneath, 1963, Principles of Numerical Taxonomy, W.H. Freeman and Company,
San Francisco and London.
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CHEMOTAXONOMY 11
11.1 WHAT IS CHEMOTAXONOMY?
The approach of taxonomy in which chemical features of plants are used in developing classifications
or in solving taxonomic problems is called Chemotaxonomy, Chemosystematics, Chemical Taxonomy,
Chemical Plant Taxonomy, or Plant Chemotaxonomy. Some of its basic features or evidentiary
characters include alkaloids, flavonoids, carotenoids, polysaccharides, terpenoids, fatty acids, amino
acids, aromatic compounds, etc.
The chemical evidences are useful in establishing relationships among taxa is exemplified
(Cronquist, 1981) by the (i) presence of aromatic plants in Juglandales, (ii) production of betalains
and not anthocyanins by the members of Caryophyllales, (iii) presence of highly aromatic compounds
in the members of Lamiaceae, (iv) presence of alkaloids in Solanaceae, and (v) presence of tannifer-
ous plants in Sapindaceae.
11.2 PURPOSE OF CHEMOTAXONOMY

Chemotaxonomy has been used at all levels of the hierarchy of classification starting from the rank
of ‘variety’ up to the rank of ‘division’ in plants. Moreover, chemical evidences have been used in
all the groups of the plant kingdom starting from simple organisms, such as fungi and bacteria, up
to the most highly advanced and specialised groups of angiosperms. These evidences are used in the
classification of plants with two main purposes, (i) to develop such taxonomic characters which may
improve the existing systems of plant classification, and (ii) to develop the present-day knowledge
of phylogeny or evolutionary relationships of plants.
11.3 A BRIEF HISTORY

The history of the knowledge or use of chemical characters of plants and their utility in plant clas-
sification is very old. Socrates, the great Greek philosopher, was forced to take his own life by
drinking a broth of poison hemlock, i.e. Conium maculatum of family Apiaceae or Umbelliferae.
In his Materia Medica, Dioscorides in the 1st century A.D. described several aromatic mints. But
122 Plant Taxonomy
some of the pioneer botanists, who employed chemical evidences in plant classification, are Nahemia
Grew (1673), William Withering (1785), A.P. de Candolle (1804), Abbott (1886) and Geshoff (1891).
However, the modern phase of chemotaxonomy began with the publication of the work of Abbott
(1886) in Botanical Gazzette. McNair published a series of papers on the use of chemical evidences
in solving taxonomical problems during 1917 and 1945.
The major research centre in the field of chemotaxonomy in the world is The Institut fur
Pharmakognosie of the University of Kiel, Germany.
Some of the books providing details of chemotaxonomy are by Hawkes (1968), Swain (1973),
Gibbs (1974), Bisby et al. (1980), Young and Seigler (1981), and Harborne and Turner (1984).
11.4 CHEMICAL CHARACTERS AND THEIR USE IN TAXONOMY

No suitable classification of the chemical characters and their use in taxonomy is developed so far.
Naik (1984) divided these characters into three categories, i.e. (i) directly visible characters, e.g. starch
grains, raphides, silica, gypsum, etc., (ii) chemical test characters, e.g. phenolics, betalains, oils, fats,
waxes, alkaloids, etc., and (iii) proteins. On the other hand, Jones and Luchsinger (1987) divided
the natural chemical plant products useful in taxonomy, on the basis of their molecular weight, into
two major groups, i.e. (i) low molecular weight compounds with a molecular weight of 1000 or less,
called micromolecules, e.g. amino acids, alkaloids, fatty acids, terpenoids, flavonoids, etc., and (ii)
high molecular weight compounds with a molecular weight of over 1,000, called macromolecules,
e.g. proteins, DNA, RNA, complex polysaccharides, etc. Sometimes, the term semantides is used for
the information-carrying proteins, such as DNA, RNA, etc.
In the absence of a suitable classification, a discussion of some taxonomically important chemical
compounds, along with a few examples of their systematic value, is undermentioned:
11.4.1 Flavonoids
Most widely and most effectively used compounds in chemotaxonomy are the flavonoids. These
are the phenolic glycosides consisting of two benzene rings linked together through a heterocyclic
pyrane ring. The reasons for their wide use in chemotaxonomy include their widespread distribu-
tion, chemical complexity, the relative physiological stability, quick and easy identification, and great
structural variation.
Some of the common classes of flavonoids include anthocyanidins (e.g. cyanidin, delphinidin,
etc.), flavones (e.g. apigenin, levtolin), biflavonyls (e.g. amentoflavone), flavonols (e.g. kaempferol,
quercetin), flavonones (e.g. naringenin), isoflavones (e.g. orobol), isoflavonoids (e.g. ferreirin), chal-
cone (e.g. butein) and aurone (e.g. sulphuretin).
Some of the specific examples of the use of flavonoids in chemotaxonomy are undermentioned:
1. Alston (1967) documented hybridization between the species of Baptisia by using chromato-
graphic patterns of flavonoids.
2. Dahlgren (1980, 1981) used flavonoids in clarifying certain relationship while suggesting his
famous system of angiosperm classification.
3. Psilotum contains biflavonyls, and this brings it closer to lycopods than leptosporangiate
ferns, as proposed by some workers.
Chemotaxonomy 123
4. Ellagic acid, a polyphenol, is confined to woody dicots and not found at all in monocots.
5. Analysis of leaf flavonoids of Liliaceae, Juncaceae, Cyperaceae and Gramineae (Williams,
1976) suggest that all these families have arisen from Liliaceous ancestors.
6. Williams et at. (1983) identified six ornamental species of Fuchsia on the basis of their leaf
flavonoids.
7. Belzer and Ownbey (1971) studied the flavonoid pattern of 5 species of Tragopogon of
Compositae and their Fl and F2 hybrids, and noted that all the species and their population
show different chromatographic results.
8. Isoflavone iridin is found only in the section Pogonivis of genus Iris.
9. South American species of Eucryphia of the family Eucryphiaceae may easily be distin-
guished on the basis of their flavonoid glycosides.
10. Species of Malus are classified on the basis of their flavonoids by Williams (1982).
11. Presence of five leaf flavonoids (tricin, luteolin, glycosyl flavones, bisulphate complex and
5-glucoside) in both Gramineae and Palmae, link them together.
12. Species of Spirodella, unidentifiable on morphological grounds, may be distinguished on the
basis of their flavonoid chemistry.
11.4.2 Betalains
Betalains differ from flavonoids and other phenolic compounds owing to the presence of nitrogen
in them. However, these are functionally equivalent to phenolics. Betalains are popularly known as
nitrogenous anthocyanins. They comprise of red to violet betacyanins and yellow betaxanthins. Some
of the aspects of taxonomic value of betalains are undermentioned:
1. All the ten families of the order Centrospermae (i.e. Aizoaceae, Amaranthaceae, Basellaceae,
Caryophyllaceae, Chenopodiaceae, Didieraceae, Molluginaceae, Nyctaginaceae, Phytolaccaceae
and Portulacaceae) produce betalains and are placed together in a single order Caryophyllales
in modern schemes of classification.
2. Betalains are also reported in the family Cactaceae. This family, once placed elsewhere (i.e.
in the order Cactales) in the classification schemes, should be placed under the betalains-
containing order Caryophyllales or in the older order Centrospermae.
11.4.3 Terpenoids
These are a biogenic group of diverse compounds with mevolonic acid as their precursor (Goodwin,
1970). Some regard them as components of essential oils. According to the number of isoprene
units present in a terpenoid molecule, they may be of five categories: (i) hemiterpenes (C5H8) e.g.
tiglic acid, (ii) monoterpenes (C10H16). e.g. myrcene, (iii) sesquiterpenoids (C15H24), e.g. parnesol,
(iv) diterpenoids (C20H32), e.g. phytol and, (v) triterpenoids (C30H48), e.g. squalene.
Terpenoids have been used extensively in the chemotaxonomy of mints, umbellifers, Citrus plants,
and gymnosperms. Some of their chemotaxonomic applications are mentioned below:
1. Origin of certain Citrus cultivars was determined by studying their rind and leaf terpenoid
pattern.
124 Plant Taxonomy
2. Geographical races of Pseudotsuga menziesii were distinguished by studying the terpenoids

of their cortical oleoresins.
3. In the taxonomy of Compositae and tribe Genisteae of Fabaceae, the petal carotenoids, which
form a prominent group of terpenoids, prove very useful.
4. Mirov (1961) used the gum terpentines in the taxonomy of Pinus.
5. Some specific terpenoids are found in certain families, e.g. cucurbitacins in Cucurbitaceae,
asperuloside in Rubiaceae, and sesquiterpene lactones in Compositae.
11.4.4 Steroids
True steroids possess two methyl groups and are mostly alcohols or esters. They are located in plant
cutins and perform the function of water proofing. Their distribution proved helpful in the taxonomy
of the genera of tribe Veratreae of family Liliaceae. These genera contain the steroid veratum accord-
ing to Kupchan et al. (1961).
11.4.5 Iridoid Compounds

Monoterpenoid cyclopentanoid lectones represent a separate class of compounds of taxonomic sig-
nificance, called iridoids. Some of the aspects of their taxonomic value are mentioned below:
1. Because of the presence of an iridoid compound, aucubin, genus Buddleia was transferred
from Loganiaceae to Buddleiaceae, near Scrophulariaceae.
2. The iridoid compound, arsperuloside, is the characteristic of Rubiaceae whereas aucubin is
found in Orobanchaceae, Scrophulariaceae and Cornaceae.
3. On the basis of the presence of iridoids, the interfamilial relationships were clarified between
Cornales, Scrophulariales, Caprifoliales, and Rubiales.
4. Several iridoid-families (e.g. Rubiaceae, Scrophulariaceae, Cornaceae, Dipsacaceae and
Caprifoliaceae) are considered the putative progenitor of Compositae.
11.4.6 Alkaloids
These are the basic nitrogen-containing organic compounds, usually with a heterocyclic ring.
Alkaloids show significant effects on the central nervous system of animals, and are well-known for
their medicinal, chemical, poisonous, and systematic viewpoints. Some of their taxonomic applica-
tions are undermentioned:
1. Jones and Luchsinger (1987) stated that alkaloids are useful in taxonomic studies in Veratrum
(Liliaceae), Papaver and Argemone of Papaveraceae, as well as in Lycopodium, Lupinus, and
Caryophyllales.
2. Morphine is produced only by Papaver somniferum, and strychnine is found only in
Strychnos.
3. Because of the accumulation of isoquinoline alkaloids containing protopine, in both
Fumariaceae and Papaveraceae, the two are considered closely related.
4. Lupin alkaloids are the characteristic of the tribes Sophoreae, Genisteae and Podalyrieae of
subfamily Lotoideae of Leguminosae.
Chemotaxonomy 125
5. Distribution patterns of alkaloids in two genera (Genista and Adenocarpus) of the tribe
Genisteae of Fabaceae and the genus Ammodendron of the tribe Sophereae of the same
family suggest the transfer of Ammodendron from Sophereae to the tribe Genisteae.
6. Several modern taxonomists, including Takhtajan, Cronquist, Thorne and Dahlgren, rear-
ranged the families with apocarpous gynoecia in Magnoliidae on the basis of the distribution
of benzylisoquinoline alkaloids.
11.4.7 Glucosinolates
Glucosinolates is a group of about 85 thioglucosides which on hydrolysis form glucose and a cor-
responding isothiocyanate. They are also called mustard oil glucosides because they are confined to
the taxa of the mustard family (Cruciferae) and some related families of Capparales.
Formerly, Cruciferae, Capparaceae, Fumariaceae and Papaveraceae were all included under one
order Rhoeadales. But the recent findings support the view that glucosinolate-producing families
(Cruciferae and Capparaceae) should be placed separately under Capparales, and the alkaloid-
containing families (Fumariaceae and Papaveraceae) under Papaverales. Members of Bataceae and
Gyrostemonaceae also contain glucosinolates. Glucosinate patterns are also used to document hybrid-
ization in some genera of Cruciferae, e.g. Cakile.
11.4.8 Amino Acids

These are the building blocks of proteins and are universally distributed in plant tissues. Several
workers have studied the taxonomic potential of amino acids. A few examples of their taxonomic
significance are mentioned below:
1. Lathyrus species could be grouped under seven infrageneric groups on the basis of the asso-
ciation of amino acids within the seeds.
2. According to Turner and Harborne (1967), canavanine is found only in the taxa of the sub-
family Lotoideae of Leguminosae.
3. Lathyrine is found only in the genus Lathyrus while azetidine-2-carboxylic acid is found
only in Agavaceae, Liliaceae and Amaryllidaceae.
4. Acetyl ornithine is identified as the main amino acid of Fumarioideae.
5. Species of the section Gummiferae of the genus Acacia can be recognised easily on the basis
of the amino acid contents of their seeds.
11.4.9 Lipids and Waxes

Lipids are the esters of fatty acids with glycerol, mostly made up of carbon, hydrogen and oxygen.
Plant waxes are esters of long-chain alcohols with long-chain fatty acids. They may contain free
alcohols, free fatty acids, aldehydes, ketones or hydrocarbons. Some of the taxonomic applications
of lipids and waxes are undermentioned:
1. Restricted distribution of some fatty acids is characteristic of certain families, e.g. petrose-
linic acid of Umbelliferae, erucic acid of Cruciferae, chaulmoogric acid of some taxa of
Flacourtiaceae, and xymenyric acid of Santalaceae.
126 Plant Taxonomy
2. Proportions of fatty acids in lipids are constant in both genera and species in Palmae (Hilditch,
1952).
3. Malvalic acid exists in the members of Malvaceae while capric acid is characteristic of
Lauraceae and Lythraceae.
4. Fatty acids of bacterial lipids are not found in other plants.
11.4.10 Nucleic Acids

Unavailability of the necessary expertise and equipment, required for the nucleic acid technique, to
most of the taxonomists, hindered the progress in this direction. However, eight species of Atriplex
of Chenopodiaceae were compared on the basis of their DNA nucleotide sequences. DNA of wheat,
rye and barley of Triticeae, compared with that of oat of Aveneae, showed marked differences.
Moreover, wheat DNA is more similar to the DNA of rye than that of barley. According to Goldberg
et al. (1972), all the 13 species of Cucurbita contain mainband and satellite DNA whose densities
are constant throughout in all species of this genus.
11.4.11 Proteins
Proteins are used in chemotaxonomy owing to their large complex molecules, universal distribution,
presence in large quantity in some organisms, and also because of the knowledge of several simple
and rapid methods of protein analysis, such as electrophoresis. Phylogenetic affinities in Triticinae
were demonstrated by Johnson and Hall (1965) by the process of protein electrophoresis. This
process also helped in establishing a close relationship between Vicia and Lathyrus. By exploiting
peroxidase, esterase and acid phosphatase pattern, Symenoidis and Tsekos (1984) suggested that the
genus Taeniantherum, formerly considered as a part of the genus Hordeum, should be treated as an
independent genus.
As the proteins represent the direct products of the DNA code, their comparative analysis may
be used as fundamental taxonomic characters. Protein analysis by electrophoresis and other methods
may be used to determine whether differences actually exist among plants that are treated as distinct
species.
11.4.12 Cyanogenic Compounds

The ability of some plants to release poisonous compounds, such as hydrocyanic acid, amygdalin,
etc., after injury to their cells, is called cyanogenesis, and these poisonous compounds are called
cyanogenic compounds. Plants containing these compounds are called cyanophoric plants. Over two
thousand vascular plants have so far been established as cyanophoric. Taxa belonging to Araceae,
Gramineae, Juncaceae, Juncaginaceae and Scheuchzeriaceae are commonly cyanogenic. Several
dicotyledonous families belonging to Asteridae, Rosidae and Dilleniidae are also cyanogenic.

1. What do you mean by chemotaxonomy? Explain its purpose and history in brief.
2. With the help of suitable examples, explain the use of chemical characters in solving taxo-
nomical problems.
Chemotaxonomy 127
3. Explain some aspects of taxonomic value of flavonoids, terpenoids, alkaloids and amino
acids.
4. Write an essay on chemotaxonomy.
Suggested Reading
Alston, R.E. and B.L. Turner, 1963, Biochemical Systematics, Prentice-Hall, Englewood Cliffs, N.J.
Bisby, F.A., J.G. Vaughan and C.A. Wright (eds.), 1980, Chemosystematics: Principles and Practices,
Academic Press, London.
Crawford, D.J. and D.E. Giannasi, 1982, Plant chemosystematics, Bioscience 32: 114–118.
Cronquist, A., 1980, Chemistry in plant taxonomy: An assessment of where we stand, (In) F.A. Bisby et
al. Chemosystematics: Principles and Practices, Academic Press, London.
Gibbs, R.D., 1974, Chemotaxonomy of Flowering Plants, 4 Vols. McGill-Queens Univ. Press, Montreal.
Harborne, J.B., 1964, Biochemistry of Phenolic Compounds, Academic Press, London.
________ 1967, Comparative Biochemistry of the Flavonoids, Academic Press, London.
________ and B.L. Turner, 1984, Plant Chemosystematics, Academic Press, New York.
Hawkes, J.G. (ed), 1968, Chemotaxonomy and Serotaxonomy, Academic Press, London.
Jones, S.B. and A.E. Luchsinger, 1987, Plant Systematics, McGraw-Hill, New York.
Smith, P.M., 1976, The Chemotaxonomy of Plants, Edward Arnold, London.
Swain, T. (ed.), 1973. Chemistry in Evolution and Systematics, Butterworth, London.
Young, D.A. and D.J. Seigler, 1981, Phytochemistry and Angiosperm Phylogeny, Praeger, New York.
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MOLECULAR
TAXONOMY 12
12.1 WHAT IS MOLECULAR TAXONOMY?
Molecular taxonomy, also known as molecular systematics, is the use of the structure of molecules
to obtain information on an organism’s evolutionary relationships. (As is well-known, the molecule
is the smallest unit of an element or compound which occurs naturally. Molecules consist of more
than one atom, e.g., a molecule of hydrogen consists of two hydrogen atoms (H2)). Some prefer to
call molecular systematics as molecular phylogenetics. The results of a molecular phylogenetic analy-
sis are expressed in the form of a tree called phylogenetic tree (Hillis and Moritz, 1996, Wikstro,
et al., 2001).
In an International Workshop on Molecular Systematics, organised at Bhagalpur University,
Bhagalpur (India) during 11–16 February 2007, Pandey (2007) outlined that “Molecular systematics
encompasses a series of approaches in which phylogenetic relationships are inferred using information
from macromolecules of the organisms under study. The type of molecular data acquired include
those from DNA sequences, DNA restriction sites, microsatellites, RAPDs (Random Amplified
Polymorphic DNA) and AFLPs (Amplified Fragment Length Polymorphism).”
12.2 BASICS USED IN THE TECHNIQUES AND APPLICATIONS OF

MOLECULAR TAXONOMY
All living organisms contain DNA, RNA, and proteins. Genetically, it has been proved that closely
related organisms generally have a high degree of agreement in the molecular structure of these
substances. On the other hand, molecules of distantly related organisms usually show a pattern of
dissimilarity. Molecular taxonomists use such data to build a “relationship tree” that provides a
picture of probable evolution of various organisms. This is actually the basic used in the techniques
and applications of molecular taxonomy.
In majority of the molecular taxonomical studies, the most common approach is the compari-
son of sequences for genes using techniques of sequence alignment to identify similarity among
organisms.
Molecular phylogeny is also applied in DNA barcoding, where the species of an individual
organism is identified using small sections of mitochondrial DNA.
Molecular Taxonomy 129
In the field of human genetics, molecular phylogeny techniques are used in (i) genetic testing to
determine a child’s paternity, and also (ii) in the branch of criminal forensics focussed on evidence
known as “genetic fingerprinting”.
12.3 NAMES OF SOME TECHNIQUES USED IN MOLECULAR TAXONOMY

Some techniques used in molecular taxonomy by the taxonomists are (i) extraction, (ii) amplification,
(iii) sequencing of genes, (iv) sequence alignment, and (v) phylogenetic analysis. Young taxonomists
should be trained in these and other related techniques by the experts by organising workshops, time
to time, in different parts of the world.
An old technique, which has now been superseded by the above-mentioned recent techniques,
was to determine the divergences between the genotypes of individuals by DNA-DNA hybridisation.
This technique was based on entire genotype rather than on particular sections of DNA.
12.4 RELATION OF CHEMOTAXONOMY AND MOLECULAR TAXONOMY

Use of chemical evidence in taxonomy is called chemotaxonomy. It makes use of proteins, enzymes,
carbohydrates and other molecules, which get separated using techniques such as chromatography.
During last few decades these techniques have been largely replaced by DNA-sequencing under
molecular taxonomy. The DNA-sequencing produces the exact sequences of nucleotides or bases in
either DNA or RNA segments extracted using different techniques. (As is well-known, nucleotide
is a molecule with a pentose sugar, a phosphate group and a purine or pyrimidine base containing
nitrogen. Nucleotides are the units which form the long chain polymers, nucleic acids i.e. DNA and
RNA. Base is a purine or pyrimidine unit).
Molecular taxonomy techniques like DNA-sequencing are generally considered superior for evo-
lutionary studies because the actions of evolution are finally reflected in genetic sequences.
12.5 WHERE DO WE STAND TODAY IN KNOWING DNA SEQUENCING

TECHNIQUES AND WHY?
It is still a long and expensive process to sequence the entire DNA of an organism, i.e., its genome.
(The genome is the genetic material on the sets of chromosomes in a cell. The smallest genome
consists of all genes on a haploid set of chromosomes). Due to this, the process of DNA-sequencing
has so far been done for only a few species.
12.6 STATUS OF MOLECULAR TAXONOMY IN INDIA

Most of the so-far globally completed molecular systematic studies have been done in the devel-
oped countries, mainly because they are long and quite expensive. However, with the rich biologi-
cal resources in many developing countries, molecular taxonomy will greatly advance the field of
systematics and several related branches “if more scientists and students in these countries are
involved in molecular systematic studies”. “India is one of the richest countries of the world as far
as biodiversity is concerned. However, not many molecular studies have been carried out due to
130 Plant Taxonomy
lack of proper training in the field of molecular systematics” (Pandey, 2007). The workshop, aim-
ing to provide training to young scientists, organised under the auspices of Indian Association for
Angiosperm Taxonomy (IAAT) at Bhagalpur University in 2007 “was the first molecular systematics
workshop on flowering plants organised in India” (Pandey, 2007).
12.7 WHAT DOES A TYPICAL MOLECULAR SYSTEMATIC ANALYSIS REQUIRE?

A typical molecular systematic analysis requires the sequencing of approximately 1000 base pairs.
At any location within such a sequence, the bases found in a given position may vary between dif-
ferent organisms. The particular sequence present in a given organism is referred to as its haplotype.
(A haplotype is actually a particular set of alleles at several very closely linked loci). Technically,
since there are four base types, with 1000 base pairs, we could have 41000 distinct haplotypes. In
a molecular systematic analysis, the haplotypes are actually determined for a well-defined area of
genetic material. At the end of the process, the haplotypes from a smaller number of individuals
from a definitely different taxon are determined. These are technically called out group. Then the
base sequences for the haplotypes are compared. Initially, the difference between two haplotypes
is determined by counting the number of locations where they have different bases. This is techni-
cally called as the number of substitutions. The difference between organisms is mentioned as the
percentage divergence. It is calculated by dividing the number of substitutions by the number of
analysed base pairs.
After the determination of the divergences between all pairs of samples, a triangular matrix of
differences is resulted. It is submitted to some definite statistical calculations, and finally a dendro-
gram is resulted. (A dendrogram is a branching diagram after the style of a family tree reflecting
similarities or affinities of some sort). From the dendrograms, it is calculated whether or not they
provide any clues about the current ideas about taxonomy of the group. Evolutionary trees are pre-
pared using all these studies.
12.8 ASSUMPTIONS AND USES OF MOLECULAR TAXONOMY

Some of the characteristics of molecular taxonomy and the assumptions involved there in are listed
below:
1. Being essentially a cladistic approach, the molecular taxonomy assumes that classification
must correspond to phylogenetic descent, and that all valid taxa must be monophyletic. (The
word cladistics refers to the method of classifying organisms into groups or taxa based on
recency of common descent, as judged by the possession of shared-derived characteristics.
A branching diagram showing the relationships between groups of organisms determined by
the methods of cladistics is called a cladogram.)
2. Molecular taxonomy often uses the molecular clock assumption which emphasizes that quan-
titative similarity of genotype is a sufficient measure of the recency of genetic divergence.
But this assumption does not always hold good in relation to speciation (evolutionary process
in which new species are produced).
3. Mitochondrial DNA is very conveniently used globally for molecular systematic analysis in
animals.
4. Molecular systematics helps in establishing the relationship of different plant groups at DNA
level.
5. It has also unlocked the treasure chest of information on evolutionary history of
organisms.
6. By the use of conserved molecular sequences, it is now possible to define and identify a very
large number of species of plants and animals.
7. Evolutionary patterns of biodiversity are also now investigated using DNA data.
8. DNA taxonomy has its definite role in genome mapping and biodiversity conservation.
9. Molecular systematics is now highly useful not only in plant identification but also in know-
ing genealogical history of plants as well as animals.
10. It also helps in understanding the phytogeography, which can ultimately help in conservation
of biodiversity.
11. DNA-based molecular markers, used for designing DNA-based molecular probes, have also
been developed under the branch of molecular systematics (details discussed elsewhere in
this chapter under Article 12.10).
12. Modern tools and techniques of DNA analysis can find out differences at single-nucleotide
level and sequences with these differences are then said to possess single nucleotide poly-
morphism (SNP). SNP studies, if carefully performed, can differentiate between two similar-
looking organisms.
13. Amplified Fragment Length Polymorphism (AFLP) technique produces large sets of poly-
morphic markers that are used to analyse closely related taxa. AFLP data are also used in
studying polyploidy evolution.
14. Molecular systematics is now used in Gene Bank search using definite methodology. With
the help of computers, sequence alignment is now done using different softwares.
15. By the techniques of PAUP (Phylogenetic Analysis Using Parsimony) cladistic analysis is
now done using both morphological and molecular data.
12.9 CRITICISM AND FUTURE OF MOLECULAR SYSTEMATICS

Above-mentioned assumptions and uses of molecular systematics are not totally uncontroversial
among biological systematists. Being a cladistic method, molecular systematics is criticised by
systematists as the science of “cladistics” in general. Some even go upto the extent of calling it a
“mistake to replace a classification based on visible and ecologically relevant characteristics by one
based on genetic details that may not even be expressed in the phenotype”.
Inspite of all this, molecular systematics is gaining increasing acceptance. As studies are proceeding
making gene sequencing easier and cheaper, the science of molecular systematics is being used and
applied for more and more groups. Considering the overall scenario, some molecular systematic
studies have already lead to radical revisions of many accepted taxonomic groups.
132 Plant Taxonomy
12.10 A NOTE ON MOLECULAR MARKERS

Molecular markers are DNA-based markers used widely in various fields including DNA-fingerprinting
and also as forensic tools. These have revolutionised the entire scenario of biological sciences and
occupied their definite position in various fields, including genetic engineering, taxonomy, physiology,
embryology, etc. They are widely used globally to bring about automation in the process of genome
analysis. The discovery of polymerase chain reaction (PCR) has brought about a new class of DNA
profiling markers which has resulted in the development of (i) marker-based gene tags, (ii) maping-
based cloning of agronomically important genes, (iii) variability studies, (iv) phylogenetic analysis,
(v) synteny mapping, (vi) marker-assisted selection of desirable genotypes, etc. (Joshi, et al., 1999).
DNA-markers can now provide data that can be analysed objectively.
Molecular markers actually include biochemical constituents (e.g., secondary metabolites in plants)
and macromolecules, viz., proteins and DNA. DNA-based molecular markers are widely used in
biological fields, and in this regard DNA-fingerprinting was introduced for the first time by Jeffrey
et al., in 1985. According to these workers, DNA-fingerprinting describes the “bar-code-like DNA
fragment patterns generated by multilocus probes after electrophoretic separation of genomic DNA
fragments” (Joshi, et al., 1999). This technique is now widely used in analysis of genome evolution,
taxonomy, plant breeding, population genetics, and diagnostics. Joshi et al. (1999) listed some prop-
erties desirable for ideal DNA-markers. These include (i) highly polymorphic nature, (ii) frequent
occurrence in genome, (iii) easy access or easy availability, (iv) easy and fast assay, and (v) high
reproducibility, etc.
Various types of molecular markers used to estimate DNA polymorphism are generally categor-
ised as (i) hybridisation-based markers, and (ii) PCR (polymerate chain reaction)-based markers.
The versatile technique of PCR was invented by Saki et al., (1985). Joshi et al., (1999) described in
detail various types of DNA-markers and their application in plant genome analysis and breeding.
One of the most recent applications of molecular markers has been shown in sex identification of
dioecious plants.
12.11 SOME SPECIFIC EXAMPLES SHOWING ROLE OF

MOLECULAR SYSTEMATICS IN PLANTS
(1) Endress (2002) reviewed the details of the morphology and angiosperm taxonomy in molecu-
lar era and concluded that the goals of comparative morphology have shifted in the pres-
ent molecular era. According to him, the “morphology no longer plays the primary role in
phylogenetic studies”. New opportunities for morphology are now opening up “that were not
present in the premolecular era”, e.g., (i) phylogenetic studies with combined molecular and
morphological analysis, (ii) reconstruction of evolution of morphological features based on
molecularly derived cladograms, (iii) redefined analysis of morphological features induced
by inconsistencies of previous molecular and molecular phylogenetic analyses, and (iv) explo-
ration of the evolution of morphological traits by integration of comparative structural and
molecular developmental genetic aspects. He has also mentioned that the field of angiosperm
taxonomy in the molecular era “is still in its infancy in botany; its advancement is one of
the major goals of evolutionary botany” (Endress, 2002).
(2) Recently, Hussain et al. (2008) studied molecular analysis of dicot-monocot split and rela-
tionship among major angiosperm groups. Using RAPD (random amplified polymorphic
DNA)-technique, they established relationship among major angiospermic groups based on
cladistic analyses. “The phylogenetic trees of relationship derived from molecular data con-
firm dicots as the ancestral class of monocots, there seems no dicot-monocot split. Dicots
form an ancestral class of magnoliids and the monocot lineage was derived from one of the
basal magnoliids. Their studies also “pinpoint dicots as the possible close relatives of the
monocots” (Hussain et al., 2008).

1. Define molecular taxonomy. What are the basics used in the techniques of molecular
taxonomy?
2. What is DNA-barcoding?
3. What is the relation of chemotaxonomy and molecular taxonomy?
4. Write a note on the status of molecular taxonomy in India.
5. Define the terms (a) DNA-sequencing, (b) dendrogram, (c) cladistics.
6. Describe the typical requirements for a molecular systematic analysis.
7. Give an account of some assumptions and uses of molecular taxonomy.
8. Write a detailed note on molecular markers.
Suggested Reading
Endress, P.K., 2002, Morphology and angiosperm systematics in the molecular era. Bot. Rev. 68:
545–570.
Hillis, D.M. and C. Moritz, 1996, Molecular Systematics (2nd ed.), Sinauer Associates Incorporated
(ISBN 0–87893–282–8).
Hussain, M.B., V. Verma and Z.A. Malik, 2008, Molecular analysis of dicot-monocot split and relationship
among major angiosperm groups. Afr. Jour of Pl. Sci. 2(1): 1–4.
Jeffrey, A.J., V. Wilson and S.L. Thein, 1985. DNA-fingerprinting. Nature 314: 67–73.
Joshi, S.P., P.K. Ranjekar and V.S. Gupta, 1999, Molecular markers in plant genome analysis. Curr. Sci.
77: 230–240.
Pandey, A.K., 2007, Molecular systematics (Report of International Workshop), Curr. Sci. 92(7):
881–882.
Saki, R.K., S. Scharf, F. Faloona, K.B. Mullis, G.T. Horn, H.A. Erlich and N. Arnheim, 1985. PCR-
technique. Science 230: 1350–1354.
Wikstro, M.N., V. Savolainen and M. Chase, 2001, Evolution of the angiosperms: Calibrating the family
tree. Proc. Roy. Soc. Lond. B. 268: 2211–2220.
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SEROTAXONOMY 13
13.1 WHAT IS SEROTAXONOMY?
The application or utility of serology in solving taxonomic problems is called serotaxonomy. The
study of antigen-antibody reactions is called serology. The substance capable of stimulating the
formation of an antibody is called antigen. A highly specific protein molecule produced by plasma
cells in the immune system is called antibody. The antibodies combine chemically with specific
kinds of antigens. Proteins are the useful carriers of taxonomic information and are most widely
used antigens in serotaxonomy.
Serology is useful in the following ways:
1. It expresses similarities and dissimilarities amongst different taxa, and these data are helpful
in taxonomy. It determines the degree of similarity between species, genera, families, etc. by
comparing the reactions of antigens from various plant taxa with antibodies raised against
the antigen of a given taxon.
2. It helps in comparing non-morphological characteristics, the knowledge of which is useful
in taxonomy.
3. Single proteins from different plant taxa are also compared by serological techniques.
13.2 SOME OTHER RELATED TERMS

1. Serology According to Boyden (1964) it is the branch of biology which deals with “the
nature and interactions of antigenic material and antibodies”. But according to Smith (1976)
serology is the “study of origin and properties of antisera.”
2. Agglutinogens Antigens are also called agglutinogens.
3. Agglutinins Antibodies that participate in agglutination process.
4. Agglutination A type of antigen-antibody reaction that results in visible clumps of organ-
isms or other materials.
Serotaxonomy 135
5. Precipitins These are the antibodies “capable of combining with and reacting upon anti-
genic material because of certain determinant groups” (Fairbrothers, 1968). Precipitins are
the antibodies causing precipitation.
6. Precipitin Reaction It is the in vitro reaction between solutions or suspensions of antigen
and antibody, with the formation of a precipitate.
7. Reference Reaction It is the “reaction between an antiserum and the antigenic material used
to stimulate its formation” (Fairbrothers, 1968).
8. Haptens These are the substances which are unable to induce antibody formation by them-
selves, but are able to react specifically with antibodies.
9. Serological Reactions Antigen-antibody reactions in vitro are known as serological
reactions.
13.3 GENERAL FEATURES OF SEROLOGICAL REACTIONS

1. The reaction is specific, an antigen combining only with its homologous antibody and vice
versa.
2. Entire molecules react and not the fragments.
3. Antigens or antibodies do not pass through denaturation during these reactions.
4. Combination occurs at the surface, and it is firm but reversible.
5. Both antigens and antibodies take part in the formation of precipitates or agglutinates.
6. Antigens and antibodies can combine in varying proportions.
13.4 BRIEF HISTORY OF SEROTAXONOMY

Nuttal (1901) was the first biologist to compare the immunochemical specificity of serum proteins
for systematic purposes. Other notable serologists of that time were Kowarski (1901), Bertarelli
(1902) and Magnus (1908). It was Dunbar (1910) who showed that proteins from pollen, seeds and
leaves of rice were serologically distinct. A school of serology was founded in 1914 by Gohlke at
Koenigsberg in Germany, and in the later years Germany became the centre of serological studies.
Mez (1926) and Moritz (1934) of Germany and Boyden (1942) of America developed the technique
of serology in the later years. Notable contributions on the application of serology to taxonomy are
those of Rives (1923), Hawkes and Lester (1966), Smith (1968, 1969, 1972), Kloz (1971), Fairbrothers
(1967, 1975, 1977, 1978, 1983) and Crawford and Giannasi (1982).
13.5 GENERAL PROCESS OF SEROTAXONOMY

In serotaxonomical studies, the protein extract of plant or animal origin, i.e. antigen, is injected into
the blood stream of a rabbit or an experimental animal, to form the antibodies. In response to a
specific antigen a specific antibody is produced. The serum, called antiserum, is then made to react
in vitro with the antigenic protein as well as with proteins of other taxa, the affinities of which are
to be determined. The amount of the precipitation shows the degree of the protein homology. The
whole process may be explained as under:
136 Plant Taxonomy
Suppose, we are to ascertain the closeness of a taxon P with taxa R, S and T. The proteins
extracted from P are injected into a rabbit, in which the antibodies are produced. The antibodies
are then extracted from the blood of the rabbit in the form of an antiserum. When the antiserum
is allowed to react with the original protein extract from P, a total coagulation takes place. When
this antiserum is allowed to react with the protein extracts from the taxa R, S and T, the degree of
coagulation varies. This degree of coagulation is related directly to the closeness between the taxa
being compared.
13.6 EXAMPLES OF SEROTAXONOMIC IMPORTANCE

Out of the several cases of the implications of serological data in the classification of angiosperms,
some are mentioned below:
1. According to Fairbrothers (1983) serological data have been used in the classification of orders
and the assignment of families in Apiales, Caryophyllales, Capparales, Fagales, Cornales,
Magnoliales, Juglandales, Papaverales, Rubiales, Ranunculales, Scrophulariales, Typhales,
Primulales, etc.
2. Fairbrothers and Johnson (1959) separated six species of Bromus on the basis of serological
studies.
3. According to Jensen (1967), serological characteristics within Ranunculaceae show a close
similarity between Aconitum-Delphinium, Actaea-Cimicifuga, Anemone-Clematis and
Ranunculus-Myosurus, and suggest a common ancestry for Aquilegia, Leptopyrum and
Thalictrum.
4. While studying the serological assessment of relationships within Solanaceae, Hawkes
and Tucker (1968) observed a strong serological relationship between Solanum, Nicotiana,
Hyoscyamus, Datura and Salpiglossis.
5. Hillebrand and Fairbrothers (1970) made a phytoserological systematic survey of Caprifoliaceae
and suggested the monophyletic origin of the family. According to these workers there is a
close relationship between Caprifoliaceae and Cornaceae on serological grounds.
6. On the basis of serological studies, Johnson and Fairbrothers (1965) suggested that the genera
Magnolia and Michelia show the closest affinity within Magnoliaceae.
7. Simon (1971) showed a close relationship between Nymphaeaceae and Nelumbonaceae on
the serological ground.
8. Serotaxonomic findings of Pickering and Fairbrothers (1970) in Umbelliferae support the
classification of the family into Apioideae, Saniculoideae and Hydrocotyloideae, and also
suggest that Apioideae is more close to Saniculoideae than to Hydrocotyloideae.
9. Gartner (1978) suggested some serological evidences for assigning Phaseolus aureus and P.
mungo to the genus Vigna.
10. Hydrastis of Berberidaceae has more serological similarities with Ranunculaceae than
Berberidaceae.
11. On the basis of serological results, Mez and Ziegenspeck (1926) prepared a stammbaum or
family tree for the entire plant kingdom.
Serotaxonomy 137
12. According to Lee and Fairbrothers (1978), two genera (Galium and Asperula) of Rubiaceae
show similar serological characters, and both of them differ from other genera of this
family.
13. Kloz (1971) applied the serological techniques in the classification of Leguminosae.
14. Hawkes and Lester (1968) showed a relationship amongst the tuber-bearing taxa of Solanum
by serological techniques.
15. Fairbrothers et al. (1975) supported the separation of Illicium from Magnoliaceae to
Illiciaceae, and of Schizandra from Magnoliaceae to Schizandraceae, on the basis of sero-
logical studies.
16. Piechura and Fairbrothers (1983), on the basis of their serotaxonomic studies, supported the
view that taxa of Oleoideae evolved from those of Jasminoideae.

1. Differentiate between serology and serotaxonomy.
2. Define the following terms in one sentence only.
(a) Agglutinogens, (b) Agglutinins, (c) Agglutination
3. Describe some ways of usefulness of serology.
4. Discuss, in brief, the general process of serotaxonomy.
5. Give at least five examples which show the importance of serotaxonomy.
Suggested Reading
Boyden, A., 1964, Perspectives in systematic serology, In C.A. Leone (Ed.) Taxonomic Biochemistry and
Serology, Ronald Press, New York, pp. 75–99.
Fairbrothers, D.E., 1977, Perspectives in plant serotaxonomy, Ann. Mo. Bot. Gard. 64: 147–160.
_________ 1983, Evidence from nucleic acid and protein chemistry, in particular serology, in angiosperm
classification, Nord. J. Bot. 3: 35–41.
Hawkes, J.G. (ed.), 1968, Chemotaxonomy and Serotaxonomy, Academic Press, London.
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EVOLUTION OF
ANGIOSPERMS 14
14.1 WHAT IS PHYLOGENY?
The history of development of a race, or simply evolutionary history, is called phylogeny.
Radford (1986) defined the word ‘phylogeny’ as “the study of the genealogy and the evolution-
ary history of groups of organisms”, and the word genealogy he defined as “the study of ancestral
relationships and lineages”. However, according to Jones and Luchsinger (1987) “the evolutionary
development or lineage of a taxon is its phylogeny”.
The phylogeny of angiosperms, in the light of the above-mentioned definitions, therefore, includes
the knowledge of the ancestors of angiosperms, place and time of their origin, the primitive angio-
sperms and the interrelationships among their major groups.
14.2 WHAT ARE ANGIOSPERMS?

Angiosperms, generally known as flowering plants, constitute that subdivision of seed plants whose
members have a stem with xylem tissue containing vessels and bear their seeds within one or more
closed carpels. Another subdivision of seed plants is gymnosperms. Seeds are enclosed within the
carpel or carpels in angiosperms while they are borne exposed on the carpels in gymnosperms.
One remarkable feature of the angiosperms is the possession of a complex reproductive structure
called flower. A flower is a structure usually containing sepals, petals, stamens and carpels. However,
there exist flowers which consist of only a single stamen or a single ovary.
Some of the major characteristics of the angiosperms are (1) vessels in the xylem, (2) phloem
contains sieve elements and companion cells, (3) embryo sac is generally Polygonum-type or eight-
nucleate (1 egg, 2 synergids, 2 polar nuclei, and 3 antipodals), (4) double fertilisation, (5) closed car-
pels, (6) after fertilization the ovary becomes a fruit, enclosing one or more seeds, and (7) a reduced
gametophytic generation. However, there exist numerous exceptions to all these characteristics.
Angiosperms have the greatest number of plant species (over 300,000 species) inhabiting the earth.
They include annuals, perennials, herbs, shrubs, and trees, ranging from very small to gigantic sizes.
Phylogeny: Origin and Evolution of Angiosperms 139
There exist parasitic, epiphytic, saprophytic and even insectivorous species among angiosperms. They
include two classes, Monocotyledoneae and Dicotyledoneae. Some of the major differences between
the monocots and dicots are undermentioned in Table 14.1:
Table 14.1 Major differences between dicots and monocots
S.No. Dicots Monocots
1. Cotyledons 2 (rarely 1, 3, or 4). Cotyledon 1.

2. Leaves mostly with reticulate venation. Leaves mostly with parallel venation.
3. Intrafascicular cambium usually present. Intrafascicular cambium absent.
4. Vascular bundles usually in rings. Vascular bundles usually scattered.
5. Floral parts usually in sets of 5. Floral parts usually in sets of 3.
6. Pollen typically triaperturate. Pollen uniaperturate.
14.3 GENERAL PRINCIPLES OF ANGIOSPERM PHYLOGENY

According to Thorne (1976), the botanists working on angiosperm phylogeny face three major prob-
lems, viz. (1) incomplete and inadequate fossil records, (2) prevalence of convergent evolution, and
(3) vast modifications in the structure of these plants. However, in spite of the meagre fossil records
of angiosperms, their available fossil data and some other characters indicate definite trends in their
phylogeny. The problem of the convergent evolution has been tackled, to some extent, by assembling
and evaluating the information from all possible sources.
On the basis of the studies of the comparative morphology and palaeobotany, Bessey (1915) pre-
sented some guiding principles of the angiosperm phylogeny. These principles were called by Bessey
as dicta. In the light of modern discoveries, these Besseyan Principles were modified by several
modern phylogenists, including Cronquist (1968), Takhtajan (1969), Hutchinson (1973), Thorne
(1976), etc. Undermentioned are some of the modified “Besseyan Principles”, which are commonly
used in the angiosperm phylogeny:
1. Usually the woody plants have preceded the herbs, climbers and vines.
2. Annuals have been derived from both biennials and perennials, and biennials have developed
from perennials.
3. Terrestrial habit preceded the aquatic or epiphytic habit.
4. Dicotyledons are more primitive than monocotyledons.
5. Alternate leaves are primitive while opposite or whorled leaves are advanced.
6. Simple leaves preceded compound leaves.
7. Bisexual flowers are primitive while unisexual flowers are advanced.
8. Multi-membered flowers are primitive while those with a few members are advanced.
9. Actinomorphic flowers preceded zygomorphic flowers.
10. Free perianth parts are primitive compared to fused perianth parts.
11. Flowers with petals preceded apetalous flowers.
12. Many free stamens are primitive compared to a few or fused stamens.
140 Plant Taxonomy
13. Hypogynous condition is primitive while epigynous and perigynous conditions are derived.
14. Apocarpous condition is primitive to the syncarpous condition.
15. Pollens having one pore preceded those with 2 or more pores.
16. Arillate seeds with a small embryo and well-developed endosperm are primitive than non-
arillate seeds with a large embryo and ill-developed endosperm.
17. Axile placentation is primitive while free-central placentation is derived.
18. Single fruits preceded aggregate fruits.
19. Capsule fruits preceded drupe or berry fruits.
14.4 ANGIOSPERM PHYLOGENY: A GENERALLY ACCEPTED PICTURE

The available fossil evidences show that the angiosperms originated roughly 165 million years ago
in the Mesozoic era, probably in the Jurassic period or perhaps even earlier than that, and appeared
in great outburst in Upper Cretaceous time i.e., about 110 million years ago. But the actual origin
or true relationships of this large group of plants, dominating the earth today, is still not known.
Earliest known fossil angiosperms are the members of Ranales as well as of Amentiferae. Because
of this, some phylogenists believe that Ranales is the basal group in the evolution of angiosperms
while others believe that Amentiferae should be considered as most primitive. But till today, all these
are mere assumptions about the possible ancestors of angiosperms. Some assumptions consider the
seed ferns to be the ancient while others consider some group of the Cycads, etc. The sum total of
all we know about it is that suddenly, in Upper Cretaceous time, there appeared a large outburst of
already highly evolved and advanced angiosperms.
14.5 MONOPHYLETIC OR POLYPHYLETIC ORIGIN

The question of phyly of the angiosperms is still not solved, mainly because of their inadequate fossil
records. But this much is sure that the angiosperms are a natural group, and contain such characters
which make them unique from all other vascular plants (Takhtajan, 1969; Stebbins, 1974).
Faegri (1980), Dahlgren (1983) and several other phylogenists believe that angiosperms are mono-
phyletic, i.e. a group originated from a single ancestor at a single time in the past. They believe
so because present-day angiosperms show remarkable consistency in their characters, i.e. uniform
staminal structure, characteristic endothecial layer of the anther wall, double fertilization, formation
of triploid endosperm, and presence of sieve tubes in all. Hickey and Doyle (1977) also support the
monophyletic origin of angiosperms on the basis of their studies of monosulcate pollens. Dahlgren
(1983) believes that the ancestor of the present-day angiosperms was a gymnospermous member.
However, no definite fossil evidences are available in favour of the monophyletic origin.
Eames (1961), Cronquist (1965), Meeuse (1970, 1975), Krassilov (1977) and several other phylo-
genists have been of the view that angiosperms are polyphyletic, i.e. dicots and monocots originated
at different times from different primitive stocks, and attained their present status through parallel or
convergent evolution. Fossil records, variety in perianth and the nature of carpel in both dicots and
monocots also support the theory of polyphylesis. The primitive orders of both the monocots and
dicots do not show any close relationship in their characters, and this also favours the polyphyletic
origin.
However, in the light of the available fossil records and other characters, it appears that angio-
sperms, as a group, are monophyletic, and their families or groups of families are polyphyletic. To
find out the exact ancestors of angiosperms, more fossil records should be investigated.
14.6 FOSSIL RECORDS OR TIME OF ANGIOSPERMS ORIGIN

The available fossil records indicate that the angiosperms spread over the entire earth from the equa-
tor to the polar regions about 100 million years ago and during a period of only about 10 million
years. During this period, the vegetation was dominated by the gymnosperms and ferns. But some
phylogenists, such as Wolf et al. (1975), believe that they originated in the early Mesozoic or even
in the late Palaeozoic, i.e. about 250 million or more years ago. However, the majority of the fossil
records point out their origin to be about 130 to 135 million years ago during the early Cretaceous,
and the same view is supported by Taylor (1981) on the basis of his studies of fossil pollens.
The angiosperms dominated the earth during the later parts of the Cretaceous period. Fossils of
the monosulcate pollen grains are reported from the lower Cretaceous rocks. Such type of pollen
grains are characteristic of the primary monocotyledons and dicotyledons, both. In the early years
of Cretaceous, the angiosperms split into monocots and dicots and both of them became prevalent
in the terrestrial flora along with ferns and gymnosperms in the later years.
Professor David Dilcher of Indiana University and his associates (Dilcher, 1979; Retallack and
Dilcher, 1981) provided sufficient evidence of the importance of fossil records in the early history
of angiosperms. They discovered a 95 million years old fossil bisexual flower, thereby indicating
the coexistence of unisexual and bisexual flowers in the early history of the flowering plants.
Fossil pollens and leaves of several Magnoliales, Ranunculales, Theales, Hammamelidales and
some monocots have been reported from the Maestrichtian stage of the Upper Cretaceous period,
i.e. about 70 million years ago. Muller (1970) reported pollen of members of Compositae from the
Oligocene of Tertiary period of the Cenozoic era, i.e. only about 25 to 30 million years ago.
14.7 PROBABLE ANCESTORS OF ANGIOSPERMS: SOME THEORIES

To identify the ancestors of the flowering plants is not easy and this aspect has been discussed
in detail by phylogenists. It is difficult to find a group of plants, that was not at sometime or
the other considered the ancestor of angiosperms. Various orders of gymnosperms (Bennettitales,
Cordaitales, Cycadofilicales, Cycadales, and Gnetales), pteridophytes (Psilophytales, Lycopodiales
and Equisetales), and even algae have been considered as the probable ancestors of angiosperms by
different workers. Charles Darwin called the origin of angiosperms an “abominable mystery” in the
absence of definite evidence which could help in tracing their probable ancestors. In spite of the
accumulation of huge literature the problem still remains unresolved.
Following are the views of various workers:
1. Isoetes-Monocotyledon Theory Campbell (1928) advocated that monocotyledons have been
derived from Isoetes via Ophioglossaceae. Linear leaves and cormous habit of Isoetes were
correlated with some simple aquatic plants of monocots. Details of the embryo of Isoetes
142 Plant Taxonomy
and the anatomy of its older sporophyte show similarity with some lower aquatic monocots,
such as Najas flexilis. But neither any fossil evidence nor similarity between the angiosperm
flower and sporophyll of Isoetes exist to support this theory.
2. Pteridosperm Theory Majority of the phylogenists (Andrew, 1947; Arnold, 1949; Thomas,
1955; Cronquist, 1968) believe that seed ferns (Pteridosperms or Cycadofilicales) were the
ancestors of angiosperms. Vascular histology, stelar structure and available fossil records also
favour this theory. The most interesting evidence in favour of the pteridosperm theory is the
absence of vessels in the secondary wood of some primitive angiosperms and in seed ferns.
In both, the pteridosperms and angiosperms, the sporangial development is eusporangiate,
and the amphiphloic stele is also common. But it is not possible to explain the complex
pteridosperm seed in terms of the simple angiosperm ovule.
Cronquist (1968), however, mentioned that although “most students of phylogeny provisionally
accept the seed ferns as the probable ancestors of the angiosperms”, yet it should be accepted
only after a definite fossil evidence is traced.
3. Caytonialean Theory Caytoniales, a Middle Jurassic order of fossil gymnosperms, seem to
have a few angiosperm-like features, and are looked upon as their possible ancestors. The
ovules in this group of seed ferns were semi-enclosed in small pouches, their leaves had
reticulate venation, and their sporangia resembled superficially with the anthers of angio-
sperms. However, the caytonialian order was strongly pinnately veined and the angiosperm
carpel is palmate. Most angiospermic ovules have two integuments while Caytoniales had
single integument.
This theory was first proposed by Thomas (1925, 1936) and was supported by Stebbins (1974)
on the basis of the homology of ovules of angiosperms with those of Caytoniales.
4. Glossopteridalean Theory Dahlgren (1983) has considered the seed fern order Glossopteridales
as the probable ancestors of angiosperms. He suggested that the “Cupules” in the members of
this order correspond to the outer integument of an angiospermic ovule. Magasporophylls of
glossopterids are identical with a typical leaf of angiosperms. However, there is no similarity
between the pollen of the two.
5. Bennettitalean Theory On the basis of the nature and organization of the reproductive struc-
tures of the members of Bennettitales and their similarity with the flower of angiosperms,
Bennettitales were considered as the ancestors of angiosperms. This theory was first proposed
by Saporta and Marion (1885) followed by Arber and Parkin (1907).
The strobili of the Mesozoic genus Cycadeoidea resemble the flowers of Magnolia. Both these
structures are bisexual and contain an elongated axis having bracts, microsporophylls and
megasporophylls. However, the stamens of Magnolia are free while the microsporophylls of
Bennettitales are connate. The seeds of Magnolia and other primitive angiosperms are with
copious endosperm and small embryo while those of Bennettitales were non-endospermic with
a large embryo. The angiospermic stem has a small pith, thick vascular cylinder and thin
cortex while that of Bennettitales had a large pith, thin vascular cylinder and thick cortex.
6. Coniferales-Amentiferae Theory Coniferales are treated as the probable ancestors of angio-
sperms by several phylogenists including Engler (1892) and Rendle (1904). They pointed
out several resemblances between conifers and angiosperms, and treated Amentiferae as the
most primitive dicots. Flowers in amentifers (Casuarinaceae, Salicaceae, Fagaceae, etc.) are
simple and naked like those of conifers. Fertilization in conifers is similar to that of angio-
sperms (Doyle, 1945). The seed-scale complex of Juniperus also resembles the gynoecia of
Amentiferae. However, definite differences prevail between the ovuliferous scales of conifers
and the angiosperm carpel.
7. Gnetales-Angiosperm Theory Gnetales, the transitional group between angiosperms and
gymnosperms, are also considered as the ancestors of angiosperms. Gnetum bears angio-
sperm-like leaves, female gametophyte and vessel-bearing wood. Gnetales have two cotyle-
dons like dicots. Their ovules bear two integuments as is the case with most of the angio-
sperms. Stamens of all the three genera of Gnetales (Gnetum, Ephedra and Welwitschia)
are similar to those of angiosperms. Gametophytes of Gnetum and Welwitschia are highly
reduced like those of angiosperms. However, the vessels of Gnetales originated in an entirely
different manner from those of angiosperms (Cheadle, 1953). Moreover, the details of the
vascular anatomy strongly refutes the suggestion of the existence of any similarity between
the two groups.
8. Durian Theory Corner (1949) believed that Durio zibethinus, a member of Bombacaceae
found in Burmese and Malayan forests, is the only “surviving member of primitive angio-
sperms”, and after the name of the genus, the theory is named as “durian-theory”. D.
zibethinus is a cauliflorous tree with large spiny capsules and fleshy arillate seeds. Arillate
genera are found in about 45 families of angiosperms according to Corner (1949), mostly
distributed in the tropics, like Durio. According to him within these arillate families, a
reduction series can be traced from genera with arillate fruits to genera having fruits of other
types. Parkin (1953) and Eames (1961) criticised this theory.
9. Pentoxylon Theory Meeuse (1961) compared Pandanus, a monocot, with Pentoxylales, a
group of fossil pteridophytes, and listed a number of common features between the two. He
proposed that Pandanus “is a direct descendent of a group of plants practically identical”
with Pentoxylales. Pandanus and Pentoxylales, according to Meeuse (1961), resemble each
other in their stem and leaf anatomy, male and female inflorescence, and pollen. Both bear
terminal tufts of strap-like leaves in spiral arrangement, copious endosperm and minute
embryo. However, Pant and Kidwai (1971) attributed these similarities, between Pandanus
and Pentoxylales, to the parallel evolution.
10. Czekanowskiales Theory Jones and Luchsinger (1987) stated that “the Czekanowskiales,
an order of Jurassic seed plants belonging to the class Ginkgopsida, are regarded by some
as early angiosperms”. They further mentioned that similar to Caytoniales, the ovule in this
order also “was surrounded by a cupule having a flange that somewhat resembled a stigmatic
surface”.
14.8 PRIMITIVE ANGIOSPERMS

Undermentioned are some of the views regarding the type of primitive living flowering plants:
1. Some phylogenists believe that the first flowering plants were upland plants that diversified in
tropical or arid regions about 250 million years ago in Permian. Absence of fossils from this
period is explained by pointing out that upland plants are usually not preserved as fossils.
144 Plant Taxonomy
2. A few other phylogenists are of the view that the primitive flowering plants were tropical
mesophytic trees with pinnate leaves and the fruits in the form of clusters of large arillate
follicles.
3. Another view is that the early flowering plants were insect-pollinated, Magnolia-like, ever-
green trees of tropical upland regions. Fossils, resembling the present-day Magnoliaceae are
common among the fossil remains of Lower Cretaceous period. Phylogenists believe that the
primitive angiosperms had a minute embryo embedded in the copious oily endosperm, and
the same is true of the embryo and endosperm of Magnolia. All these characters led some
botanists to call Magnoliales as Living Fossils.
4. According to Stebbins (1974), primitive angiosperms were small woody plants with small
leaves and moderate-sized flowers.
5. According to the Takhtajan (1980), the primitive angiosperms were woody plants with
simple, entire, pinnately-veined leaves; moderate-sized, solitary, axilary or terminal flower
having perianth of modified bracts; leaf-like stamens; monosulcate pollen; and conduplicate
carpel.
6. Hutchinson (1973) believed that the primitive angiospermic flowers were medium-sized
and grouped together in lateral clusters, much like those of Drimys winteri of the family
Winteraceae.
7. Engler and Prantl (1925) and Wettstein (1935) regarded Amentiferae as the primitive angio-
sperms because their flowers do not possess petals, and inflorescences resemble the cones
of gymnosperms.
8. Bailey (1956) regarded Ranales as the most primitive order of living dicotyledons. This is
supported by the evidences from floral anatomy, wood anatomy and pollen grain structure.
14.9 PLACE OF THE ORIGIN OF ANGIOSPERMS

Where the angiosperms might have first originated, is also not yet finally decided. Three divergent
views regarding the place of their origin are undermentioned:
1. Heer (1868) suggested that the angiosperms originated at high latitudes in the north polar
regions. Hooker, Asa Gray, Arnold, etc. supported his view. From the polar Arctic and
Antarctic regions the angiosperms spread in successive waves across the whole earth. However,
Croizat (1952) and Takhtajan (1969) rejected firmly the Arctic or Antarctic origin of flower-
ing plants.
2. Hallier (1912) proposed that the angiosperms originated first in the basin of the Pacific Ocean,
at places such as New Zealand, New Caledonia, Andes and the Hawaiian Islands. According
to Takhtajan (1969), the western part of the basin of Pacific Ocean is particularly rich in the
primitive angiospermic families and covers the areas such as Assam, Burma, China, Japan
to Australia, New Zealand, New Caledonia and Fiji, and therefore the angiosperms probably
originated in this part of the eastern and south-eastern Asia and Australia.
3. According to Dutoit (1947), the discovery of some angiosperm-like inflorescences from
the Triassic beds of South Africa indicates that this must be the place of the origin of
angiosperms.
14.10 ORIGIN OF MONOCOTS: SOME VIEWS

1. Hegelmaier (1878) suggested that the embryo of monocots originated as one of the two
cotyledons of a typical dicot embryo that failed to develop. This hypothesis was named as
abortion hypothesis. This view was suppored by workers such as Metcalfe (1936) and Eames
(1961).
2. Engler (1892) proposed that monocots originated from Isoetales, Ophioglossales and a few
other pteridophytes.
3. Sargant (1903, 1908) proposed that monocots originated as a result of the adaptation of the
aquatic habit to the geophyllous or marshy habit, and called them as aquatic geophytes.
4. Hallier (1914) concluded that monocots originated from a dicotyledonous family
Lardizabalaceae.
5. Cheadle (1953) has shown that vessels in dicots and monocots originated independently,
and therefore the ancestors of monocots must have been some vesselless dicots such as
Nymphaeales or Magnoliales.
6. Hutchinson (1959) opined that monocots originated from Ranunculaceae.
7. Takhtajan (1961) believed that monocots might have originated from some extinct ves-
selless herbs having apocarpous gynoecium and monocolpate pollen similar to the modern
Nymphaeales.
8. Stebbins (1974) was of the opinion that monocots probably originated from some Drymis-like
ancestor having vesselless wood and short internodes.
9. Burger (1977) opined that monocots were derived from some dicot ancestors similar to
Chloranthaceae of Piperales. He believed that Piperales were related more closely to monocots
than any other living dicots. He suggested that simple flowers, like those of Chloranthaceae,
came together by loosing their internodes and formed trimerous flowers of several monocots
and Piperales.
14.11 LINES OF EVOLUTION IN ANGIOSPERMS

Ranales are considered as the most primitive angiosperms because of (i) absence of vessels in over
100 living representatives of families (Magnoliaceae, Winteraceae, Alismaceae, etc.) of this order,
(ii) presence of a triradiate ridge on the pollen grains of the genus Schizandra of Magnoliaceae,
(iii) binucleate pollen at the time of its release from a microsporangium in several Magnoliaceae,
Winteraceae, Ranunculaceae, etc., and (iv) peculiar floral morphology and anatomy, etc.
The general opinion of phylogenists is that at least three main lines of general evolution are
observed from the Ranalian plexus. (1) The first line consists of Rhoeadales, Sarraceniales, Parietales,
Malvales, Geraniales, Sapindales, Rhamnales and some Tubiferales, (2) the second line attains the
climax of Campanulales through Rosales, Myrtales, Umbellales and Rubiales, and (3) the third line
included Liliales and Orchidales.
In the first line the hypogynous condition remained unchanged. Rhoeadales and Sarraceniales
evolved parallaly from Ranales. Development of the gynophore in Capparidaceae shows a definite
relation between Parietales and Rhoeadales. Malvales evolved through Bombacaceae. There exist
146 Plant Taxonomy
definite relationships between Malvales and Geraniales. Geraniales are related to Sapindales, and
Sapindales to Rhamnales. Fusion of sepals, petals and carpels, as well as the epipetalous condition is
seen in Solanaceae. Labiatae is the most advanced family of this line. Besides the fusion of sepals,
petals and carpels and epipetalous condition, the number of floral parts in Labiatae are reduced to
four or two.
In the second line of evolution a change is observed from hypogyny to epigyny through perigyny.
The change from hypogyny to epigyny is seen clearly in Rosales. The formation of hypanthodium
shows a change of perigyny into epigyny in Myrtales. Epigyny and syncarpy in Umbellales show
an advancement over Myrtales. Umbellales are related to Rubiales through the characters such as
umbel-like inflorescence, suppression of calyx, epigyny, etc. Campanulales top this line with the
presence of pappus, irregular flowers, and fusion of floral parts.
The third line shows a few resemblances between monocotyledons (e.g. Alismataceae) and dicoty-
ledons (e.g. Ranunculaceae). Liliales and Ranales differ in the number of floral parts and fusion of
carpels. Epigynous condition of Iridaceae and Amaryllidaceae is a stage of advancement over Liliales.
There exist evidences of the origin of Gramineae from Liliaceae. But the reduction in number of
stamens and carpels and also the loss of perianth parts indicate that Gramineae are more advanced
than Liliaceae. Orchidaceae possess insect-pollinated, irregular and showy flowers, and is thus the
most advanced family of monocotyledons.

1. What do you mean by the word ‘phylogeny’? Explain some general principles of angiosperm
phylogeny.
2. Write brief notes on:
(a) Monophyletic and polypyhyletic origin of angiosperms.
(b) Primitive angiosperms.
3. Describe at least five theories, put forward by phylogenists, regarding the “probable ancestors
of angiosperms”.
4. Explain some views of scientists about the place of origin of angiosperms.
5. Phylogenists have opined at least three different lines of evolution of angiosperms. Explain
them briefly.
6. Discuss some basic aspects of the origin and evolution of angiosperms.
Suggested Reading
Axelrod, D.I., 1952, A theory of angiosperm evolution, Evolution. 6: 29–60.
Burger, W.C., 1981, The monocot theory of angiosperm evolution, Evol. Theory 5: 189–225.
Cheadle, V.I., 1953, Independent origin of vessels in monocotyledons and dicotyledons, Phytomorphology
3: 23–44.
Daghlian, C.P., 1981, A review of the fossil record of monocotyledons, Bot. Rev. 47: 517–555.
Doyle, J.A., 1978, Origin of angiosperms, Ann. Rev. Ecol. Syst. 9: 365–392.
Krassilov, V.A., 1977, The origin of angiosperms, Bot. Rev. 43: 13–176.
Meeuse, A.D.J., 1975, Origin of the Angiosperms—Problem of inaptitude, Phytomorphology 25: 373–379.
Pant, D.D. and P.F. Kidwai, 1971, The origin and evolution of flowering plants, J. Indian Bot. Soc. 50A:
242–274.
Puri, V., 1967, The origin and evolution of angiosperms, J. Indian Bot. Soc. 46: 1–14.
Scott, R.A., E.S. Berghoorn and E.B. Leopold, 1960. How old are the angiosperms? Am. J. Sci. 258A:
284–299.
Stebbins, G.L., 1974, Flowering Plants: Evolution Above the Species Level, The Belknap Press,
Cambridge.
Takhtajan, A., 1969, Flowering Plants: Origin & Evolution, Smithsonian Inst. Press, Washington.
Tutin, T.G., 1952, Phylogeny of flowering plants: Facts or fiction? Nature (London) 169: 126–127.
Wolfe, J.A., J.A. Doyle and V.M. Page, 1975, The basis of angiosperm phylogeny: Palaeobotany, Ann. Mo.
Bot. Gard. 62: 801–824.
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BOTANICAL
LIBRARY 15
15.1 BOTANICAL LIBRARY AND SYSTEMATICS
A botanical library is a storehouse of botanical books, journals, periodicals, magazines, photographs,
movies, newspapers, dissertations, etc. All printed materials, drawings, films, paintings, and other
similar articles belonging to the plant sciences are also stored there in a proper system.
It is also a documentation centre of botanical activities, and stores the data related to all the
branches of plant sciences. It may also be called as a service facility to the botanists as it provides
the required literature.
A botanical library exists for the benefit of the entire society in general and for the persons inter-
ested in botany in particular. However, it is such a huge repository of the world’s botanical literature
that no single individual can utilize this system in its entirety.
The important taxonomic publications, available in the taxonomy section of a botanical library
include floras, monographs, literature indexes, and name indexes. The information regarding the
plants of a given geographical region are available in a flora, while a monograph contains informa-
tion on the systematics of a given taxonomic group of plants on a worldwide basis. The binomial
system of nomenclature is used in both the floras and monographs. Various systems of indexing
and abstracting the taxonomic literature are available in Biological Abstracts and Bibliography of
Agriculture. For the name indexes of plants, the world botanists rely on books such as Index Kewensis,
Gray Herbarium Index and J.C. Willis’s A Dictionary of the Flowering Plants and Ferns.
15.2 COMMON TERMS USED IN A BOTANICAL LIBRARY

1. Document A written item such as a letter, a manuscript, notes, an article or a book is called
a document.
2. Documentation An introduction to the written documents, specimens, etc. used as an evi-
dence or a source of information in taxonomic studies is called documentation.
3. Literature Retrieval It is the process of locating and identifying by referring to the books,
manuals, articles, and the exact information within the entire published literature that is of
interest to the user.
Botanical Library 149
4. Document Retrieval It is the process of locating and identifying both the published as well
as unpublished documents such as specimens, manuscripts, illustrations, notes, films, etc.
5. Voucher It is a specimen or written document used for the verification of information.
6. Citation A quotation from a book or author is called a citation.
7. Reference A source of information with a definite direction is called a reference.
8. Reference Book An authoritative book consulted for facts or the background information is
called a reference book.
9. Bibliography The list of books, research articles, or other writings, usually of one author
or on one subject or topic is called bibliography.
15.3 USE OF BOTANICAL LIBRARY

Radford (1986) mentioned that the botanical library is useful to the systematists as an information
source, a documentation centre, a data storehouse, and also as a service facility.
1. From the botanical library, taxonomists procure information to verify facts about the available
literature, procedure, citation, classification, and all other related aspects of his/her project,
and in that way the library is used as an information source.
2. The library is used as a documentation centre as it is a repository of rare publications, first
editions, original manuscripts, illustrations, photographs, etc.
3. The library is also used as a data storehouse for research because it stores all the published
data related to the systematic projects.
4. The big libraries are used as a service facility for providing the technical know-how of their
available resources. A reference service of these libraries helps in procuring the requisite
material, photocopies, microfilms, necessary for the projects in systematics. Most of the big
libraries also have an interlibrary loan service for supply of materials not available locally.
15.4 BOTANICAL LIBRARY AS A TRAINING CENTRE

Several big libraries train the students for using the library in a proper and systematic manner, and
thus function as training centres. The trained library staff trains the young students by (i) introduc-
ing them the references of common use, such as manuals, dictionaries, atlases, maps, encyclopedias,
indexes, etc., (ii) familiarising them with the literature of the current developments such as periodi-
cals, journals, monographs, recent bibliographies, etc., (iii) informing them of the proper rules for
making catalogues, bibliography, etc., and (iv) providing them the way of analyzing the literature
for scientific writings, etc.
15.5 CLASSIFICATION USED IN A BOTANICAL LIBRARY

In the botanical libraries, two systems of classification are used, i.e. the Dewey Decimal (DD)
system and the Library of Congress (LC) system. The Dewey Decimal system is an old system of
classification and is now being replaced by the LC-system in majority of the libraries of the world.
For details of these systems, readers may refer Swift (1970).
150 Plant Taxonomy
15.6 MAJOR BOTANICAL LIBRARIES OF THE WORLD

A huge taxonomic literature is available in five great national libraries of the world, viz. Library of
Congress in Washington, U.S.A.; Library of the Academy of Sciences in Leningrad, Russia; The
National Library of Beijing, China; Library of the British Museum in London, England; and The
Bibliotheque Nationale in Paris, France.
The other big American university libraries are at California (Berkeley), Cornell, Harvard, Iowa
State, Minnesota, Illinois, Columbia, Wisconsin, Chicago, Michigan, Yale and Texas.
Several large botanical libraries are associated with the botanical gardens, museums and botanical
societies. Chief among them are the libraries associated with the British Museum, the New York
Botanical Garden, Missouri Botanical Garden, Gray Herbarium, Massachusetts Horticultural Society
at Boston, and Academy of Natural Sciences at Philadelphia.
Some major Indian botanical libraries are associated with The Botanical Survey of India, Kolkata;
National Botanical Garden, Lucknow; Forest Research Institute, Dehradun; Indian Agricultural
Research Institute, New Delhi; and universities of Delhi, Chennai, Meerut, Allahabad, and Kolkata.
A big botanical library was established at Meerut College, Meerut by Professor V. Puri during 1948
and 1967. Till 1970, it used to subscribe 166 research journals. It also contains over 8,000 books.
15.7 FUTURE INFORMATION SYSTEMS

The botanical libraries have been the main information systems during the last 500 years, i.e. since
the invention of the movable type. But, the introduction of the high-speed electronic computers during
the last few decades has brought about a colossus change in the entire scene. Instead of the printed
pages, a large amount of the botanical information is now stored in computers in all parts of the
world. With the help of the data stored in computers, the plants may be identified more easily and
correctly. One may say it with confidence that the computers are now the main information tools.
Detailed taxonomic data are now available on the pages of a computer memory, and these data are
now easily available to the entire world via internet. The taxonomists are now able to get the latest
information on a topic of his/her choice through computers. But this does not mean that botanical
libraries are soon going to be replaced by computers. However, the present days are now of internet,
and detailed taxonomic information is now easily available in computers.

1. How will you define a botanical library?
2. Write a note on “botanical library as a training centre”.
3. Give one major difference between Dewey Decimal (DD) system and Library of Congress
(LC) system with reference to a botanical library.
4. Give a brief account of major botanical libraries of the world.
5. With reference to a botanical library, explain the following terms.
(a) Document (b) Literature retrieval
(c) Reference book (d) Bibliography
Botanical Library 151
Suggested Reading
Swift, L.H., 1970, Botanical Bibliographies: A Guide to the Bibliographic Materials Applicable to Botany,
Burgess Publish. Comp., Minnesota.
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HERBARIUM 16
16.1 HERBARIUM AND ITS LIMITS
A collection of dried and pressed plants arranged according to a classification system and available
for study or reference is known as herbarium (plural, herbaria), a name first applied by Linnaeus.
Plant specimens are usually mounted on a sheet of high quality paper. Properly dried, pressed,
and identified plant specimens are placed in thin paper folders (species covers), which are kept
together in thicker paper folders (genus covers), and finally they are incorporated into the herbarium
cupboards in their proper position. Some fleshy members (e.g. Cactaceae) are preserved in liquid
preservatives instead of being pressed and dried. Bulky plant parts (e.g. dry fruits, cones of several
gymnosperms, etc.) are dried without pressing and are stored in special boxes.
A herbarium may contain a few hundred locally collected plant specimens kept in a small place,
or it may contain millions of them collected from different parts of the world and housed in a very
big building. Herbaria in different countries remain associated with colleges, universities, scientific
societies, research institutes, botanical gardens, or well-funded government organizations. They may
contain a local collection, or flora of a district, state, country, continent or several continents. There
are also private herbaria.
The world’s largest herbarium is at Royal Botanic Garden, Kew, Richmond, Surrey, England, con-
taining over 5 million specimens. But findings of Holmgren et al. (1981) suggest that the Herbarium
of the Museum of Natural History, Paris contains over 6.5 million plant specimens. The biggest
Indian herbarium is of Botanical Survey of India at Kolkata holding over 1.3 million specimens
(Holmgren et al. 1981).
Over 1600 world’s most important herbaria are listed in Index Herbariorum of Holmgren et al.
(1981). Missouri Botanical Garden publishes a valuable monthly newsletter, entitled Herbarium News,
covering the news of different herbaria of the world.
According to Shelter (1969), the herbarium resources of the world include as many as 250 million
specimens. However, the total number actually reported from different institutions of the world was
only 148 million specimens.
Herbarium 153
16.2 HISTORY OF HERBARIUM DEVELOPMENT

Luca Ghini (1490–1556) of Italy has been the initiator of the art of herbarium. He collected plants,
dried and affixed them on paper in the form of herbarium specimens. Gherards Cibo, a student of
Ghini, started collecting and preserving plant specimens in 1532. John Falconer, an Englishman,
learned the art of preparing a herbarium, perhaps from Luca Ghini, and prepared one as early as
1553. Holmgren et al. (1981) mentioned that the great herbarium of the Museum of Natural History,
Paris was founded in 1653.
Up to the time of Linnaeus (1707–1778), the usual practice of preparing a herbarium was mounting
the specimens and binding them into volumes. The plants were usually sewn by thread. Linnaeus
started the current practice of mounting specimens on single sheets of paper and storing them hori-
zontally. By such a simple beginning of Linnaeus, herbaria have developed into facilities of hous-
ing millions of specimens, usually in steel cases. The Herbarium of the Indian Botanical Gardens,
Kolkata (now known as Central National Herbarium) was established in 1832.
In U.S.A., the oldest herbarium was started at Salem College in 1772. The herbarium of Academy
of Sciences, Philadelphia was started in 1812, University of Michigan in 1838, Missouri Botanical
Garden in 1857, and U.S. National Herbarium in 1868. The large herbaria of U.S.A. are those of
the Harvard University, the New York Botanical Garden, and the Missouri Botanical Garden.
In the earlier days the herbaria existed only in the form of a few pressed and dry specimens
of local or regional significance. But now, many of them are the centres of advanced research in
the field of taxonomy. Herbaria of Paris, Kew, Leningrad, Cambridge, Geneva and Kolkata are big
centres of taxonomic research.
16.3 A MODERN HERBARIUM

According to Fosberg (1946), a modern herbarium is a “great filling system for information about
plants, both primary in the form of actual specimens of the plants and secondary in the form of
published information, pictures and recorded notes”. However, according to Radford (1986), a modern
herbarium “is a research, training, and service institution that serves as a reference centre, docu-
mentation facility, and data storehouse”. It includes “diverse collections of flowering plants, gym-
nosperms, ferns, mosses, liverworts, lichens, fungi, algae, and fossils.” It also contains “microscope
slides, photographs, photomicrographs, wood specimens, camera lucida drawings, field notebooks,
diaries, letters, unpublished reports, manuscripts, reprints, and botanical illustrations” according to
Radford (1986).
Modern herbaria are utilized as reference centres for identification of plants by the botanists doing
research work in taxonomy, ecology, agriculture, pharmacy, etc.
Modern herbaria are the documentation centres because they contain the specimens of new taxa,
plants belonging to new discoveries, plants of economic importance, voucher specimens of cytological
and cytogenetic studies, photographs of important plants, and all documents related with systematic
research.
Geneticists, chemists, pharmacists, etc. use the modern herbaria as a data storehouse. They store
data on ecology, habitat, distribution of plants, etc.
154 Plant Taxonomy
Modern herbaria provide training for young undergraduates, solve queries of scientists, environ-
mentalists, doctors, public school students, etc., and thus work as service institutions.
16.4 TWENTY MAJOR HERBARIA OF THE WORLD

Undermentioned is the list of 20 major herbaria of the world, along with the number of their speci-
men holdings as worked out by Holmgren et al. (1981):
1. Museum of Natural History, Paris.............................................................................6.5 million
2. Royal Botanic Gardens, Kew ................................................................................over 5 million
3. Komarov Botanical Institute, Leningrad ..............................................................over 5 million
4. Conservatory and Botanical Garden, Geneva ..............................................................5 million
5. Combined Herbaria, Harvard University, Cambridge ...............................................4.5 million
6. New York Botanical Garden, Bronx ......................................................................... 4.3 million
7. U.S. National Herbarium, Washington ...................................................................... 4.1 million
8. British Museum of Natural History, London ...............................................................4 million
9. University of Lyon, Lyon ........................................................................................... 3.8 million
10. Natural History Museum, Vienna .............................................................................. 3.5 million
11. Missouri Botanical Garden, Saint Louis ................................................................... 2.9 million
12. Field Museum of Natural History, Chicago ..............................................................2.4 million
13. University of Uppasala, Uppasala ..............................................................................2.2 million
14. National Botanical Garden of Belgium, Brussels ..................................................... 2.1 million
15. Botanical Garden and Botanical Museum, Berlin .......................................................2 million
16. Academy of Natural Sciences, Philadelphia.................................................................2 million
17. Royal Botanic Garden, Edinburgh ............................................................................. 1.7 million
18. University of California, Berkeley ............................................................................. 1.5 million
19. University of Michigan, Ann Arbor .......................................................................... 1.4 million
20. Botanical Survey of India, Kolkata ........................................................................... 1.3 million
However, the latest estimates show that in India, there exist two more herbaria having more than
a million specimens. According to Tiagi and Kshetrapal (1988) these are (1) The Central National
Herbarium, Indian Botanical Gardens, Kolkata (25,00,000 specimens), and (2) Herbarium of the
Forest Research Institute, Dehradun (15,00,000 specimens). But in majority of the other Indian
books, it is mentioned that herbarium of F.R.I. Dehradun has only about 300,000 specimens and
not 15,00,000.
16.5 MAJOR INDIAN HERBARIA

Undermentioned is the list of some major Indian herbaria, along with the approximate number of
their specimen holdings as mentioned by Naik (1984), and Tiagi and Kshetrapal (1988):
Herbarium 155
1. The Central National Herbarium, Kolkata ................................................25,00,000 specimens

2. Herbarium of Forest Research Institute, Dehradun ....................................3,00,000 specimens
(15,00,000 according to Tiagi and Kshetrapal, 1988)
3. Botanical Survey of India
(i) Eastern Circle Herbarium, Shillong.................................................................... 10,00,000
(ii) Southern Circle Herbarium, Coimbatore ...............................................................1,75,000
(iii) Western Circle Herbarium, Pune ............................................................................. 50,000
(iv) Northern Circle Herbarium, Dehradun.................................................................... 42,000
(v) Central Circle Herbarium, Allahabad ...................................................................... 45,000
4. Blatter Herbarium, Mumbai ........................................................................................... 1,00,000
5. National Botanical Garden Herbarium, Lucknow ........................................................... 80,000
6. Herbarium of Industrial Section, Indian Museum, Kolkata ............................................ 50,000
7. Herbarium of Rajasthan University, Jaipur ...................................................................... 30,000
8. Herbarium of School of Plant Morphology, Meerut College, Meerut ............................ 25,000
9. Herbarium of Delhi University, Delhi .............................................................................. 15,000
10. Herbarium of Jiwaji University, Gwalior .......................................................................... 15,000
16.6 FUNCTIONS OF HERBARIA

Some of the general functions of herbaria are mentioned below:
1. Plant specimens are permanently stored in herbaria, and therefore they are the major sources
of information about plants and vegetation.
2. Preserved specimens of herbaria are used in almost all types of taxonomic research.
3. A picture of all species of a genus, or all the genera of a family may be gathered only in
the herbarium.
4. The classification of the world’s flora is based mainly on the herbarium material.
5. List of the endangered species of any region may be prepared only by herbarium
specimens.
6. Limited individual collections are identified only with the help of herbaria.
7. Monographs of genera or families are prepared only by the herbarium specimens.
8. Our knowledge of the distribution of plants, evolution, and several taxonomic problems etc.
is based mainly on the herbarium specimens.
9. Big herbaria provide training to young students in herbarium practices.
10. Exact area, region or location of the occurrence of important plants may be gathered from
the herbarium specimens.
11. Herbaria preserve type specimens, and thus serve as a repository of chromosomes, chemot-
axonomy, and experimental voucher specimens.
156 Plant Taxonomy
12. Herbarium material is used in studying the palynology, anatomy, and chemical aspects of
desired plants.
13. Herbaria provide loan of specimens for study at other institutions.
14. Herbaria preserve the national plant wealth, and provide scientific information to the public
regarding the plants.
16.7 PRECAUTIONS FOR USING HERBARIUM

1. Handle the plant specimens with extreme care because they are of scientific value and gener-
ally irreplaceable.
2. Dry specimens are brittle and easily damaged, and so keep the specimen sheets flat.
3. Store the specimens always in herbarium cases.
4. Keep the materials in folders when not in use.
5. Never keep heavy books or other heavy objects on the specimens.
6. Support the specimens with a ventilator when carrying them even for short distances.
7. If you notice that specimens are being damaged by insects, etc. inform the Curator
immediately.
8. Do not write anything on sheets of herbarium, unless permitted to do so.
9. Do not reshelve specimens in the herbarium cases.
10. If you have taken specimens on loan from another institution, return them before the loan
period is over.
11. The specimens, borrowed from other institutions, should be properly housed in fire-proof,
dust-proof and pest-free cases.

1. What do you mean by the word “herbarium”? Name the largest herbarium of the world.
Where is it located and in which country?
2. Write a brief note on “a modern herbarium”.
3. Make a list of 5 major herbaria of the world and also 5 major Indian herbaria.
4. Write six general functions of herbaria.
5. Comment on ‘Modern herbaria work as service institutions’. What precautions does a her-
barium require?
Suggested Reading
Croat, T.B., 1978, Survey of herbarium problems, Taxon 27: 203–218.
DeWolf, G.P., 1968. Notes on making an herbarium. Arnoldia 28: 69–111.
Herbarium 157
Fosberg, F.R. and M. Sachet, 1965, Manual for tropical herbaria, Regnum Vegetabile 39: 5–132.
Holmgren, P.K., W. Keuken and E.K. Schofield, 1981, Index Herbariorum,, Part I. The Herbaria of the
World, 7th ed. Bohn, Scheltema and Holkema, Utrecht.
Shelter, S.G., 1969, The herbarium: Past, present and future, Proc. Biol. Soc. Washington 86: 687–758.
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BOTANICAL
GARDENS 17
17.1 WHAT ARE BOTANICAL GARDENS?
Botanical gardens are the institutions that maintain the living plant collections of different varieties
of plants, including the ornamental and cultivated ones, wild, medicinal, of economic importance,
of various geographical regions, of special interest, etc. They are of value not only to the botanists,
horticulturists and foresters but also to the millions of tourists.
A big botanical garden contains plant species from several corners of the globe. It also includes
greenhouses, a library, a herbarium, research laboratories, and several miscellaneous resources includ-
ing photographs, paintings, illustrations, reprints, note-books and specimens of several types. It is,
therefore, not merely a garden but a botanical institution.
Modern botanical gardens serve as centres for documentation, research, reference, data storage,
education, conservation, and several other biological facilities to mankind.
At present there are over 600 botanical gardens in the world.
17.2 45 MAJOR BOTANICAL GARDENS OF THE WORLD:

A CHRONOLOGICAL DIRECTORY
The gardens were maintained in all parts of the world since the early times also, as is evident from
the old Indian classics such as Mahabharat, Ramayan and Vedas. For food, drugs, and beautiful
flowers, man had begun to cultivate plants in gardens long before the dawn of history. The Hanging
Gardens of Babylon are considered among the wonders of the ancient world. Several kinds of fruit,
vegetable and drug plants were cultivated during the Middle Ages. Romans maintained small gardens
as sources of medicine and as aids to medical studies.
Undermentioned are the historical details of some major botanical gardens of the world in a
chronological order.
17.2.1 Botanical Gardens Established During 16th Century

1. The credit of establishing the first botanical garden of the world goes to Luca Ghini (A.D.
1490–1556), an Italian Professor of Botany. He established a botanic garden at Pisa, Italy
Botanical Gardens 159
in 1543. He was its first director followed by Andrea Cesalpino (1554–1558). This garden
does not exist today.
2. Otto Botanico, the botanical garden of University of Padua, Italy, was established by
Professor Francis Bonafede on June 29, 1545. A large herbarium and a library are attached
to this garden. More than 600 portraits of botanists are displayed in this garden.
3. University Botanical Garden at Florence, Italy was established in 1545 by Luca Ghini.
4. Vatican Garden at Rome was established in 1556 by Michele Mercati, a student of Andrea
Cesalpino.
5. Bologna Botanical Garden at Bologna, Italy was founded in 1567, and its first director was
Ulisse Aldrovandi (1567–1605).
6. Leiden University Botanical Garden was established in 1587 at Leiden, Netherlands. Carolus
Clusius, the Flemish-Austrian botanist, who was the director of this garden, was the first to
establish the first greenhouse of the world in this garden in 1599.
7. Montpellier University Botanical Garden at Montpellier, is one of the earliest botanical
gardens in France. It was founded in 1593.
8. Botanical Garden, Heidelberg, Germany was also established in 1593.

1. Botanical Garden of Strasbourg, France comes next in the chronological order. It was estab-
lished in 1619.
2. Oxford University Botanic Garden, Oxford, England was established in 1621. This was the
first botanical garden of Great Britain, and Earl of Danby was its initiator. In 1734, the first
greenhouse of England was established in this garden.
3. Jardin des Plantes, a botanical garden of Paris, was established in 1640 under the name
“Jardin du Roy”. King Louis XIII of France ordered in 1626 to establish this royal botanic
garden for the purpose of growing all kinds of medicinal herbs. It was actually an indepen-
dent scientific institution, founded by the king to promote the teaching of pharmaceutical
botany. Its present herbarium and palaeobotanical departments are among the world’s best.
It is actually the oldest and most important non-university botanical garden, existing today
in the world.
4. Botanical Garden of Groningen, Netherlands was established in 1642.
5. Botanischer Garten, one of the great botanical gardens of the world, was established in
1646 in Berlin, Germany. A.W. Eichler, Adolf Engler and L. Diels were the directors of this
garden.
6. Botanical Garden of Uppasala, Sweden was founded in 1655 by O.J. Rudbeck. Linnaeus
worked as the director of this garden from 1741 to 1777, and the original part of this garden
is now maintained as a memorial to Linnaeus. With the efforts of Linnaeus, a collection of
more 300 species of this garden grew to over 3000 species within seven years.
7. Royal Botanic Garden of Edinburg, Scotland was established in 1670. Its first director was
J. Sutherland.
8. The Chelsea Physic Garden was founded in London, England in 1673, as the garden of the
Society of Apothecaries.
160 Plant Taxonomy
9. Amsterdam Botanical Garden, Netherlands was founded in 1682.

10. Botanical Gardens of Tokyo Imperial University were founded in 1684.

1. Botanic Garden of University of Moscow, Russia, was founded in 1707.
2. Druggist’s Garden of St. Petersburg, Russia, was established in 1713. St. Petersburg is the
Leningrad of today. The reorganized form of this garden is now V.L. Komarov Botanical
Institute of the Academy of Sciences, Leningrad, Russia. The herbarium attached to this
garden is one of the largest in the world, holding over 5 million specimens.
3. First botanical garden in America was established by John Bartram in 1731 near
Philadelphia.
4. Botanical garden of Vienna, Austria, was founded in 1754.
5. Jardin Botanico de Madrid, Spain, was established in 1755.
6. Royal Botanical Garden,1 Kew, England, owes its establishment in 1760 (Radford, 1986).
However, Core (1955) stated that these gardens “were officially opened in 1841, with Sir
William J. Hooker as the first director”. This is the largest botanical garden of the world.
7. Botanic Garden of Cambridge University, Cambridge, England, was founded in 1762.
8. First economic gardens, for the cultivation of spices, etc., were established on the island of
St. Vincent, British West Indies, in 1764.
9. Hortus Botanicus of University of Budapest, was established in 1771.
10. Botanical Garden of Coimbra, Portugal, was founded in 1773.
11. Indian Botanical Garden,2 Kolkata, formerly known as Royal Botanic Gardens or Kolkata
Botanic Garden, was founded in 1787 by Lt. Colonel Robert Kyd. It is “one of the great
botanical gardens of the world” (Core, 1955).

1. The Jardin Botanico de Rio de Janeiro, Brazil, was established in 1808.
2. Universitets Botaniske Have is the oldest botanical garden of Oslo, Norway, founded in
1814.
3. First botanical gardens of Australia are the Botanic Gardens of New South Wales, Sydney.
These were founded in 1816.
4. Botanic Garden at Buitenzorg, Java, was founded in 1817. It is well known for the cultiva-
tion of coffee and rubber.
5. Conservatoire et Jardin Botaniques de Geneva were also founded in 1817 in Geneva. A.P.
de Candolle was its first director.
6. Botanical Garden of Cape Town was founded in 1848.
7. Government Botanical Garden, Kingston, Jamaica was laid out in 1857.
1
For details see Artide No. 17.5.
2
For details see Article No. 17.4.
8. Singapore Botanic Gardens were established in 1859.

9. Missouri Botanical Garden, St. Louis, is the first great garden of USA. It was established
by Henry Shaw in 1859, and named as “Shaw’s Gardens”.
10. Arnold Arboretum at Harvard University, U.S.A. was founded in 1872 by James Arnold. Its
first director was C.S. Sargent (1872–1927).
11. New York Botanical Garden in New York city was established in 1891. It is one of the great-
est gardens of the world. Its first director was N.L. Britton.
12. Jardin Botanico Municipal of Buenos Aires, Argentina is the last well-known botanical garden
of the world established during 19th century. It was founded in 1892.

Almost every university with a Botany department, now has a botanical garden. Some of the major
gardens established during the early part of the century are undermentioned:
1. Botanical Garden, National Museum of Natural History, Beijing (formerly Peking), China,
was founded in 1930.
2. The Jardin Botanique de Montreal of Canada was established in 1936.
3. Botanical Garden, Forest Research Institute, Dehradun, India was established in 1934 under
the leadership of C.E. Parkinson.
4. National Botanic Garden, Lucknow, India was established in 1946.
17.3 MAJOR BOTANICAL GARDENS OF INDIA

17.3.1 Lalbagh or The Mysore State Botanical Garden, Bangalore
This famous botanical garden is considered to be the best in South India for its layout, maintenance,
scenic beauty and scientific interest. Because of roses and other red-coloured flowers it was named
as “Lalbagh” by Hyder Ali in 1760. Major Waugh was its director during 1799–1819. He introduced a
number of foreign exotic plants in this garden. However, Dr. Cleghorn made it a real botanic garden
in 1856. A tropical nursery was established in the garden in 1908. Rao Bahadur H.C. Jayaraja was
the first Indian director of this garden. The garden is now a big centre of horticultural activities. It
now has well-equipped laboratories for seed-testing and soil-testing, and also a grape orchard, tree
nursery, fruit nursery, pot garden, economic garden, and a herbal garden.
17.3.2 Lloyd Botanic Garden, Darjeeling

It was started as a branch of the Royal Botanic Garden, Kolkata on a 40 acre land piece in
Darjeeling. The land was donated by Mr. William Lloyd. Mr. A.G. Jeffrey was the first curator of
this garden. Since 1910, this garden has become a major institution for the distribution of seeds,
bulbs, and plants of temperate Himalayas to different parts of the world. It has a vast collection
of plants from Myanmar, China and Japan. It has separate sections of coniferous and indigenous
plants. A Rock Garden, Orchidarium, Bulbous section, Succulent section, Seed section, Herbarium
of over 30,000 specimens, and Rosary are its major attractions. Its coniferous section has 45 species
including Australian Callitris.
162 Plant Taxonomy
17.3.3 National Botanic Garden, Lucknow

This famous Indian botanical institution, established in its new form in 1946, is now known as
National Botanical Research Institute, Lucknow. It is popularly known as Sikander Bagh, a name
given by Nawab Wajid Ali Shah after his beloved Begum Sikander Mahal. Sikander Bagh, originally
laid out by Nawab Sadat Ali Khan (1789–1814), was converted into a botanical garden in 1946 by
Professor K.N. Kaul, its first director. The present garden and its laboratories are spread over 27
acres of land on the bank of river Gomti. Popular attractions of this garden are its Rosarium, Palm
house, Cactus house, Fern house, Orchid house, and Orchards of mango, Citrus and guava. It has
well-equipped laboratories of Plant Morphology, Aromatics, Cytogenetics, Plant breeding, Tissue
culture, Virology, Palynology, Plant Physiology, Entomology, etc. The garden bears an added experi-
mental research station at Banthra, about 20 km from Lucknow.
17.3.4 Botanical Garden of Forest Research Institute, Dehradun

Established in 1934 under the leadership of C.E. Parkinson, F.R.I. botanical garden is now among the
famous gardens of the world. It covers an area of about 20 acres in New Forest Estate, Dehradun,
and is the main Indian centre of research in problems related with plant introduction. There are
about 700 species of plants belonging to about 400 genera and about 100 families in this garden.
Over half of these 700 species have been introduced from different parts of the world. The garden
has a greenhouse, a cactus house and a Plant Introductory Nursery. Its biggest attraction is a big
herbarium holding over 30,00,00 plant specimens from all over the world.
17.4 LARGEST BOTANICAL GARDEN OF INDIA

Indian Botanical Garden, Kolkata, is the largest and oldest botanical garden of India. Formerly
named as Royal Botanic Garden or Kolkata Botanic Garden, it was established in 1787 by Lt. Col.
Robert Kyd. He founded it “not for the purpose of collecting rare plants as things of mere curiosity
or furnishing articles for the gratification of luxury, but for establishing a stock for disseminating
such articles as may prove beneficial to the inhabitants”. The garden covers an area of about 273
acres of land. William Roxburgh, The Father of Indian Botany, was its second director and founded
the world famous herbarium of this garden. The garden is now under the control of Botanical Survey
of India. Dr. K Biswas was the first Indian to be appointed Superintendent of this garden in 1937.
The garden is now noted for potato cultivation and introduction of jute, sugarcane, tea, and quinine-
yielding Cinchona. Cultivation of Aloe, coffee, India-rubber, cardamon, and Henbane are some of
the special achievements of this garden.
The Great Banyan tree (Ficus benghalensis; Moraceae; Fig. 17.1), which is one of the largest trees
in size in the world and mentioned also in the Guinees Book of World Records, is the main centre
of attraction of this garden. It appears like a miniature forest in itself. Over 2880 of its prop roots
are actually rooted in the ground. The circumference of the canopy of this single tree is more than
404 metre. It is considered to be over 250 years of age.
There are over 15000 species of plants in this garden from several countries. Some main attrac-
tions of the garden are its Palm-house, Orchid-house, Pinetum, Fernary, Cacti-collection, the giant
water lily, Victoria regia, and the section of medicinal plants.
Fig. 17.1 “Great Banyan Tree”, the largest tree in the world, mentioned also in Guinees Book of
World Records.
17.5 LARGEST BOTANICAL GARDEN OF THE WORLD

Royal Botanical Garden, Kew, officially opened in 1841, and originally established in 1760 because
of the interest of Princess Augusta of Wales, is the largest botanical garden of world. William Aiton
was the first Curator of this garden. After the death of Princess Augusta in 1772, her Kew garden
was united with the gardens of the Palace of Richmond, and the extensive area of the two gardens
was named as Royal Botanic Garden, Kew, Richmond, England. At present, the herbarium of this
garden contains over 5 million specimens, its arboretum has over 7000 species, and its glasshouses
have over 13000 species. Its Jodrell Laboratory provides best research facilities in the world for
cytologists, anatomists, geneticists and physiologists. A beautiful Alpine house, Rose garden, Bamboo
garden, and a Lily pond are also attached to this garden. Because of all these facilities and vast
developments, Kew Gardens are called the botanical capital of the world.
Some other remarkable features of Kew are Chelsea Physic Garden and world’s best rock gar-
dens. It is also well-known for some of its famous publications such as Flora of British India, Index
Kewensis and Kew Bulletin.
After 1838, this garden was transferred from British Crown to British Parliament, and Sir William
Jackson Hooker took over as its first director in 1841.
17.6 ROLE OF BOTANICAL GARDENS

1. Botanical gardens provide information of local flora and are used in the preparation of
monographs.
2. Several gardens supply seeds and material for botanical investigations.
164 Plant Taxonomy
3. They provide information on food plants, ornamental plants, medicinal plants, etc.
4. Modern gardens supply living plant resources for research in systematics, horticulture, ecol-
ogy, genetics, etc.
5. They also provide information on the protection of endangered species, and propagation of
rare plants.
6. Big botanical gardens provide training facility to younger students by providing them labora-
tory, classroom, greenhouse, nursery, etc.
7. They supply facility for courses in local flora, horticulture, hybridization, plant propagation,
etc. Their educational programmes include workshops and training sessions for teachers,
students, naturalists, etc.
8. Gardens practice and provide training for the conservation of nature.
9. They provide instructions for home gardening, and care of plants in winter and summer.
10. Botanical gardens provide aesthetically pleasing environment and thus play a major role in
providing sound mental health.

1. “A botanical garden is not merely a garden but a botanical institution”. Explain.
2. Approximately how many botanical gardens exist today in the world?
3. Write, at least, 3 names each of the botanical gardens of the world established during the
16th century and the 17th century, along with their years of establishment.
4. Who established the first botanical garden of the world at Pisa, Italy in 1543?
5. Describe briefly any three major botanical gardens of India.
6. The largest and oldest botanical garden of India is located in the eastern part of our country.
Where is it located? Write a detailed note on this garden.
7. Describe in brief the largest botanical garden of the world.
8. Mention any ‘five’ major roles that botanical gardens play in our life.
Suggested Reading
Avery, G.S. Jr., 1957, Botanic gardens—What role today?, Amer. J. Bot. 44(3): 268–271.
Hill, A.W., 1915, The history and functions of botanic gardens, Annl. Miss. Bot. Gard. 2: 185–223.
Hyams, E. and W. Macquitty, 1969, Great Botanic Gardens of the World, Nelson, New York.
Stafleu, F.A., 1969, Botanical gardens before 1818, Boissiera 14: 31–46.
Thompson, P.A., 1972, The role of botanic garden, Taxon 21(1): 115–119.
Wyman, D., 1970, How to establish an arboretum or botanic garden? Arbort. Bot. Gard. Bull. 4(52):
52–60.
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FLORAL
FORMULA AND
FLORAL DIAGRAM 18
18.1 WHAT IS A FLORAL FORMULA?
The floral formula is the numerical representation of various parts of a flower. It also depicts the
symmetry, interrelationship of various floral parts, and unisexuality as well as bisexuality in the
flower.
Actually, the floral formula is a method of summarizing the characters of a flower by a kind of
shorthand notation that exposes the essential facts. It describes the floral morphology of angiosperms
with the aid of certain standard symbols. These symbols, when combined, constitute the so called
floral formula for a particular flower.
18.2 SYMBOLS EMPLOYED IN FLORAL FORMULAE

There has been no attempt to standardize the symbols used in floral formulae on a universal basis.
Different authors have used different symbols for the same part, e.g. calyx is variously designated as
K, C and CA, and corolla as C, CO and COZ. However, the generally accepted symbols, used in the
floral formula presentation by the taxonomists of Indian subcontinent, are presented in Table 18.1,
and the same usage has been followed in the present book.
Each letter (i.e. K, C, P, A and G) is followed by the number of parts in its particular group,
e.g. K5 means 5 free sepals in the calyx whereas K(5) means 5 fused or united sepals in the calyx.
If more than one whorl is present, each is shown separately, e.g. K 2+4 means the presence of two
whorls of calyx, of which one whorl contains 2 free sepals while the other whorl contains 4 free
sepals. The same symbols are used in the case of corolla. P3+3 means the presence of two whorls
of perianth, each of 3 free tepals, e.g. Liliaceae.
A2+4 means a tetradynamous condition of stamens, i.e. 2 stamens in one whorl and 4 stamens
in the other whorl, e.g. Cruciferae. A2 + 2 means a didynamous condition, i.e. out of the 4 stamens,
2 are long and two short, e.g. Labiatae. A(μ) stands for a monadelphous condition i.e. many stamens
united in one group, e.g. Malvaceae. A(9)+1 indicates a diadelphous condition, i.e. stamens are united
in two bundles (9 stamens in one bundle and 1 separate), e.g. Papilionaceae.
166 Plant Taxonomy
Table 18.1 Floral Symbols
Br Bracteate C A Epipetalous stamens

Ebr Ebracteate P A Epiphyllous stamens
Brl Bracteolate Std Staminodes
Epik Epicalyx G4 Tetracarpellary, free carpel
0 (zero) Absence of a particular whorl G(4) Tetracarpellary, syncarpous, ovary
μ Indefinite number superior (hypogynous)
≈ Actinomorphic G(4)– Tetracarpellary, syncarpous, ovary
Zygomorphic semi-inferior (perigynous)
Male flower G(4) Tetracarpellary, syncarpous, ovary
inferior (epigynous)
Female flower
G(4) Tetracarpellary, syncarpous, ovary
Bisexual flower, or hermaphrodite condition either superior or inferior
K Calyx, sepals Pistd. Pistillode
C Corolla, petals A G Androecium and gynoecium united
P Perianth 2+4 2 in one set and 4 in another
A Androecium 2–4 2 or 4 or 2 to 4
G Gynoecium X Variable
() Cohesion of floral parts in a whorl
An arc between two floral parts marked above them indicates their adhesion, e.g. C5, A5 indicates
the epipetalous conditions of stamens; P5, A5 indicates the epiphyllous condition of stamens and;
A5, G (2) indicates the adhesion of androecium with gynoecium.
A line above the number of carpels in the gynoecium, i.e. G(5) indicates an epigynous condition
or inferior ovary, whereas a line below the number of carpels in the gynoecium, i.e. G(5) indicates
a hypogynous condition or superior ovary. A line at the side of the number of carpels in the gynoe-
cium, i.e. G(5)—indicates a perigynous condition or semi-inferior ovary.
The floral formula of Ranunculus (buttercup) i.e. Br, ≈, , K5, C5, Aμ, Gμ indicates a bracteate
(Br), actinomorphic (≈), hermaphrodite ( ) flower with a calyx (K) of 5 free sepals, corolla (C)
of 5 free petals, androecium (A) of an indefinite number of free stamens and a gynoecium (G) of
an indefinite number of free carpels with a superior ovary. On the other hand, the floral formula
of Campanula, i.e. Br, ≈, , K5, C(5), A5, G(5) shows a bracteate, actinomorphic, hermaphrodite
flower with 5 free sepals, 5 united petals to form a gamopetalous corolla, 5 free stamens, and 5
fused carpels to form a syncarpous gynoecium with an inferior ovary.
18.3 WHAT IS A FLORAL DIAGRAM?

A diagram, illustrating the relative position and number of parts in each of the sets of organs com-
prising a flower, is called the floral diagram (Fig. 18.1).
A floral diagram visually depicts the essential features of a flower in cross section. The parts of
the flower are drawn by semidiagrammatic symbols or ideographs. The number of the whorls and
Floral Formula and Floral Diagram 167
inflorescence axis
petal
carpel
stamen
sepal
receptacle
A
B bract
inflorescence axis
stigma style stamen
stamen petal
sepal carpel ovule
ovules receptacle
sepal
C
bract
D
Fig. 18.1 A-B, Median vertical section of flower (A) and floral diagram (B) of Ranunculus; C-D, Median
vertical section of flower (C) and floral diagram (D) of Campanula.
floral parts are both illustrated in a floral diagram. This also illustrates the fusion of the floral parts
as well as the flower symmetry.
A floral diagram is thus the diagrammatic representation of the ground plan of a flower depicting
the arrangement of various floral parts as projected in the transverse plane. In brief, a floral diagram
is an ideal ground plan of a flower.
18.4 WHAT DOES A FLORAL DIAGRAM INFORM?

A floral diagram furnishes the following information:
1. Sex and symmetry of a flower.
2. Bracteate or ebracteate, and bracteolate or ebracteolate nature of a flower.
3. Number of floral cycles of a flower, i.e. its tetracyclic or pentacyclic nature.
4. Number of a floral parts in each cycle of a flower.
168 Plant Taxonomy
5. If the number of sepals or petals is odd, the floral diagram informs about the anterior or
posterior position of the odd sepal or petal.
6. Type of aestivation of sepals and petals.
7. Number of whorls of stamens, their position in relation to the petals, attachment with the
petals or their free nature, monothecous or dithecous condition, and introrse or extrorse
nature, are all shown in a floral diagram.
8. Carpel number, their free or fused nature, number of locules, number of ovules in each
locule, and the type of placentation are all shown by a floral diagram.
9. Position of all floral parts in relation with the mother axis.
18.5 HOW TO DRAW A FLORAL DIAGRAM?

The floral diagram is always drawn circular in outline. The different floral whorls are represented
in concentric circles, the sepals on the outermost circle, then the petals, the stamens and carpels
towards the inner side (Fig. 18.2).
posterior side
mother axis
diagonal plane
androecium gynoecium
lateral or
transverse
plane
corolla
calyx
anterior side
bract
median or postero-anterior plane
Fig. 18.2 Floral diagram of Catharanthus roseus showing different planes of flower.
The first step to draw a floral diagram is to examine mature floral buds which are due to open
shortly but have not yet opened. Pluck the floral bud from the mother axis only after you have noted
down the anterior and posterior sides. Floral parts are drawn in a floral diagram as they would be
seen in their transverse sections below the mother axis. Make the floral diagram in the following
sequential stages:
Floral Formula and Floral Diagram 169
1. A very small circle is drawn above the floral diagram. This circle represents the mother
axis. In actinomorphic flowers the mother axis circle is denoted as ≈, but in zygomorphic
flowers it is drawn as . If the flowers are terminal, the mother axis is not drawn.
2. In bracteate flowers, a section of bract is drawn below the floral diagram. In flowers without
any bract, such a section is not drawn.
3. In bracteolate flowers, bracteoles are drawn in section on the left and right sides of the
diagram.
4. Note the number of sepals, their arrangement in relation to the mother axis and their aes-
tivation. Draw transverse sections of sepals between the mother axis and the bract, keeping
all these points in view. In case of odd number of sepals, the odd sepal would be drawn
either posterior or anterior to the flower, i.e. opposite the mother axis or opposite the bract,
respectively.
5. The same procedure is repeated for petals as for sepals mentioned above. However, petals
should be drawn alternate with the sepals.
6. If the flower is zygomorphic, petals are drawn of unequal sizes. Same may be the case with
the sepals also in zygomorphic flowers.
7. If any sepal or petal is spurred, it is shown by drawing a loop at the back of that particular
part in the floral diagram.
8. If parts of sepals or of petals are fused, draw lines to connect their edges together in the
floral diagram.
9. In the epipetalous condition (i.e. when stamens are joined with petals), link the stamens and
petals with small radial lines.
10. In case of bilabiate calyx or corolla, the two lips are joined by bulging lines.
11. Count the number of stamens, the number of whorls in which they are arranged, their cohe-
sion and adnation to other floral parts, their position in relation to petals, their introrse or
extrorse position, and draw them inside the petals in the floral diagram. Stamens are rep-
resented through transverse sections of the anthers. In the obdiplostemonous condition, the
stamens of the outer whorl are drawn opposite to the petals. Introrse stamens face towards
the centre whereas the extrorse towards the petals. Staminodes are represented either by an
asterisk (*) or by a cross (¥).
12. The gynoecium is represented by a transverse section of the ovary. Also draw the number of
locules and the number of ovules in each locule, in the ovary. Type of placentation is also
drawn,

1. What is a floral formula? How does it differ from floral diagram?
2. Which symbols are used in a floral formula for the following?
(a) Calyx (b) Corolla
(c) Absence of a particular whorl (d) Zygomorphic
170 Plant Taxonomy
(e) Actinomorphic (f) Bisexual flower

(g) Epipetalous stamens (h) Female flower
(i) Tetracarpellary, syncarpous, superior (j) Androecium and gynoecium united.
3. Explain various sequential stages of drawing a floral diagram. Illustrate your answer by
drawing a floral diagram.
Suggested Reading
Featherly, H.I., 1954, Taxonomic Terminology of the Higher Plants, Iowa State College Press, Ames.
Rangaswamy, N.G., 1966, Floral diagrams and formulae: A reappraisal, Sci. & Cult. 31(1): 33–34.
Schaffner, J.H., 1916, A general system of floral diagrams, Ohio Jour. Sci. 16: 300–360.
Steam, W.T., 1983. Botanical Latins (3rd ed.), David and Charles, Newton Abbot, England.
POSITION OF SOME C
H
SELECTED FAMILIES A
P
IN CLASSIFICATION T
E
SYSTEMS PROPOSED R
BY BENTHAM AND
HOOKER, ENGLER
AND PRANTL,
HUTCHINSON,
19
TAKHTAJAN,
CRONQUIST, AND
THORNE
Position of some selected families in some selected and widely accepted systems of classification is
presented in Table 19.1. Majority of these families are discussed in the present book. Classification
systems proposed by Bentham and Hooker (1862–1883) in their Genera Plantarum, Engler and
Prantl (1887–1899) in 20 volumes of their Die Naturlichen Pflanzenfamilien revised by H. Melchior
and E. Werdermann (1954–1964) in the form of Syllabus der Pflanzenfamilien, Hutchinson (1973)
in his treatment entitled The Evolution of Flowering Plants, Takhtajan (1980) in Botanical Review,
Cronquist (1981) in his book entitled An Integrated System of Classification of Flowering Plants,
and Thorne (1983) in Nordic Journal of Botany, were taken into consideration in the present
comparison.
Table 19.1 Position of selected families in various systems of classification
Family Bentham and Engler and Hutchinson Takhtajan Cronquist Thorne 172
Hooker Prantl (1973) (1980) (1981) (1983)
(1862–1883) (1954–1964)
Dicotyledons Dicotyledoneae Dicotyledones Magnoliopsida Magnoliopsida Annonidae
(Dicots) (Dicots) (Dicotyledoneae)
1. Magnoliaceae Ranales Ranales Magnoliales Magnoliales Magnoliales Annonales
2. Annonaceae Ranales Ranales Annonales Annonales Magnoliales Annonales
3. Ranunculaceae Ranales Ranales Ranales Ranunculales Ranunculales Berberidales
4. Nymphaeaceae Ranales Ranales Ranales Nymphaeales Nymphaeales Nymphaeales
5. Papaveraceae Parietales Rhoeadales Rhoeadales Papaverales Papaverales Berberidales
6. Fumariaceae (Treated under (Treated under Rhoeadales Papaverales Papaverales (Treated under
Papaveraceae) Papaveraceae) Papaveraceae)
7. Capparidaceae Parietales Rhoeadales Capparales Capparales Capparales Capparales
8. Brassicaceae Parietales Rhoeadales Brassicales Capparales Capparales Capparales
(Cruciferae)
9. Violaceae Parietales Parietales Violales Violales Violales Violales
10. Caryophyllaceae Caryophyllinae Centrospermae Caryophyllales Caryophyllales Caryophyllales Chenopodiales
11. Portulacaceae Caryophyllinae Centrospermae Caryophyllales Caryophyllales Caryophyllales Chenopodiales
12. Malvaceae Malvales Malvales Malvales Malvales Malvales Malvales
13. Sterculiaceae Malvales Malvales Tiliales Malvales Malvales Malvales
14. Tiliaceae Malvales Malvales Tiliales Malvales Malvales Malvales
15. Bombacaceae Malvales Malvales Malvales Malvales Malvales Malvales
16. Oxalidaceae (Treated under Geraniales Geraniales Geraniales Geraniales Geraniales
Geraniaceae)
17. Geraniaceae Geraniales Geraniales Geraniales Geraniales Geraniales Geraniales
18. Rutaceae Geraniales Geraniales Rutales Rutales Sapindales Rutales
19. Meliaceae Geraniales Geraniales Meliales Rutales Sapindales Rutales
20. Rhamnaceae Celastrales Rhamnales Rhamnales Rhamnales Rhamnales Rhamnales
21. Vitaceae Celastrales Rhamnales Rhamnales Rhamnales Rhamnales Cornales
22. Sapindaceae Sapindales Sapindales Sapindales Sapindales Sapindales Rutales
(Acerales)
23. Anacardiaceae Sapindales Sapindales Sapindales Rutales Sapindales Rutales
Plant Taxonomy
(Contd.)
24. Leguminosae or Rosales Rosales Leguminales Fabales Fabales Rutales
Fabaceae
(including
Papilionaceae,
Caesalpiniaceae
and Mimosaceae)
25. Rosaceae Rosales Rosales Rosales Rosales Rosales Rosales
26. Combretaceae Myrtales Myrtiflorae Myrtales Myrtales Myrtales Myrtales
27. Myrtaceae Myrtales Myrtiflorae Myrtales Myrtales Myrtales Myrtales
28. Lythraceae Myrtales Myrtiflorae Myrtales Myrtales Myrtales Myrtales
29. Cucurbitaceae Passiflorales Cucurbitales Cucurbitales Cucurbitales Violales Violales
30. Cactaceae Ficoidales Opuntiales Cactales Caryophyllales Caryophyllales Chenopodiales
31. Umbelliferae Umbellales Umbelliflorae Umbellales Cornales Apiales Araliales
(Apiaceae) (Apiales) (Treated
under
Araliaceae)
32. Rubiaceae Rubiales Rubiales Rubiales Gentianales Rubiales Gentianales
33. Asteraceae Asterales Campanulatae Asterales Asterales Asterales Asterales
(Compositae)
34. Sapotaceae Ebenales Ebenales Ebenales Ebenales Ebenales Ebenales
35. Primulaceae Primulales Primulales Primulales Primulales Primulales Primulales
36. Oleaceae Gentianales Contortae Gentianales Oleales Scrophulariales Oleales
Position of Some Selected Families in Classification Systems
37. Asclepiadaceae Gentianales Contortae Gentianales Gentianales Gentianales Gentianales

(Treated under
Apocynaceae)
38. Apocynaceae Gentianales Contortae Apocynales Gentianales Gentianales Gentianales
39. Boraginaceae Polemoniales Tubiflorae Boraginales Polemoniales Lamiales Solanales
40. Convolvulaceae Polemoniales Tubiflorae Solanales Polemoniales Solanales Solanales
41. Solanaceae Polemoniales Tubiflorae Solanales Scrophulariales Solanales Solanales
42. Scrophulariaceae Personales Tubiflorae Personales Scrophulariales Scrophulariales Bignoniales
43. Bignoniaceae Personales Tubiflorae Bignoniales Scrophulariales Scrophulariales Bignoniales
44. Pedaliaceae Personales Tubiflorae Bignoniales Scrophulariales Scrophulariales Bignoniales
45. Acanthaceae Personales Tubiflorae Personales Scrophulariales Scrophulariales Bignoniales
46. Lamiaceae Lamiales Tubiflorae Lamiales Lamiales Lamiales Lamiales
(Labiatae)
173
(Contd.)
47. Verbenaceae Lamiales Tubiflorae Verbenales Lamiales Lamiales Lamiales
48. Chenopodiaceae Curvembryae Centrospermae Chenopodiales Caryophyllales Caryophyllales Chenopodiales
174
49. Amaranthaceae Curvembryae Centrospermae Chenopodiales Caryophyllales Caryophyllales Chenopodiales
50. Polygonaceae Curvembryae Polygonales Polygonales Polygonales Polygonales Polygonales
51. Nyctaginaceae Curvembryae Centrospermae Thymeleales Caryophyllales Caryophyllales Chenopodiales
52. Loranthaceae Achlamydosporae Santalales Santalales Santalales Santalales Santalales
53. Euphorbiaceae Unisexuales Geraniales Euphorbiales Euphorbiales Euphorbiales Euphorbiales
54. Urticaceae Unisexuales Urticales Urticales Urticales Urticales Urticales
55. Cannabinaceae Unisexuales Urticales Urticales Urticales Urticales Urticales
56. Moraceae (Treated under Urticales Urticales Urticales Urticales (Treated under
Urticaceae) Urticaceae)
57. Casuarinaceae Unisexuales Verticillatae Casuarinales Casuarinales Casuarinales Casuarinales
58. Salicaceae Ordines Anomali Salicales Salicales Salicales Salicales Violales
Monocotyledons Monocotyledoneae Monocotyledones Liliopsida Liliopsida Liliidae
(Monocots)
59. Orchidaceae Microspermae Microspermae Orchidales Orchidales Orchidales Liliales
60. lridaceae Epigynae Liliflorae Iridales Iridales Liliales Liliales
61. Amaryllidaceae Epigynae Liliflorae Amaryllidales Liliales (Treated under (Treated under
Liliaceae) Liliaceae)
62. Musaceae Epigynae (in Scitamineae Zingiberales Zingiberales Zingiberales Zingiberales
Scitamineae)
63. Zingiberaceae Epigynae (in Scitamineae Zingiberales Zingiberales Zingiberales Zingiberales
Scitamineae)
64. Cannaceae Epigynae (in Scitamineae Zingiberales Zingiberales Zingiberales Zingiberales
Scitamineae)
65. Liliaceae Coronarieae Liliflorae Liliales Liliales Liliales Liliales
66. Commelinaceae Coronarieae Farinosae Commelinales Commelinales Commelinales Commelinales
67. Juncaceae Calycinae Liliflorae Juncales Juncales Juncales Commelinales
68. Arecaceae Calycinae Principes Palmales Arecales Arecales Arecales
(Palmae)
69. Typhaceae Nudiflorae Pandanales Typhales Typhales Typhales Typhales
70. Araceae Nudiflorae Spathiflorae Arales Arales Arales Arales
71. Cyperaceae Glumaceae Glumiflorae Cyperales Cyperales Cyperales Commelinales
72. Poaceae Glumaceae Glumiflorae Graminales Poales Cyperales Commelinales
Plant Taxonomy
(Gramineae)
Position of Some Selected Families in Classification Systems 175

1. Under which order has Magnoliaceae been placed in the systems of classification proposed
by Bentham and Hooker, Hutchinson, and Thorne?
2. Brassicaceae (Cruciferae) has been placed under order Parietales by Bentham and Hooker,
Rhoeadales by Engler and Prantl, Brassicales by Hutchinson, and ________ by Takhtajan
(1980) and Cronquist (1981).
3. Bombacaceae has been placed under same order in almost all systems of classification. What
is the name of that order?
4. Under which order has Asteraceae been treated in almost all systems of classification?
5. What are the new names of the following families?
(i) Gramineae (ii) Palmae
(iii) Labiatae (iv) Umbelliferae
(v) Compositae
Suggested Reading
Bentham, G. and J.D. Hooker, 1862–1883, Genera Plantarum, 3 Vols., London.
Cronquist, A. 1981., An Integrated System of Classification of Flowering Plants, Columbia Univ. Press,
New York.
Engler, A. and K. Prantl, 1954–1964, Syllabus der Pflanzenfamilien, 12th ed. 2 Vols., H. Melchior and
E. Werdermann (eds.), Gebruder Borntraeger, Berlin.
Hutchinson, J., 1973, The Families of Flowering Plants, 3rd ed., Clarendon, Oxford.
Takhtajan, A., 1980, Outline of the classification of flowering plants (Magnoliophyta), Bot. Rev. 46:
225–359.
Thorne, R.F., 1983, The classification of angiosperms (Annonopsida), Nordic J. Bot., 3: 102–111.
C
H
A
P
T
E
R
550 TERMS
OF PLANT
DESCRIPTION 20
20.1 PLANT PARTS1
1. Bud An undeveloped, vegetative or floral shoot, covered with protective scales, or consisting
of a short axis bearing primordia of leaves or floral parts.
2. Flower Reproductive structure of angiosperms, consisting usually of sepals, petals, stamens,
and carpels. After fertilization the ovules of flower develop into seeds.
3. Fruit Mature ovary of flowering plants containing the seeds.
4. Leaf Produced from the buds on the stem, the leaves are photosynthetic and transpir-
ing organs of the plant. They are usually green and expanded, and have a wide range of
forms.
5. Root Organ of a plant that develops initially from the radicle, grows down into the soil,
and functions for absorption and anchorage.
6. Seed Fertilized ripe ovule of flowering plants.
7. Stem Organ of a plant that develops initially from the epicotyl, grows mostly above the
ground, and functions for support and conduction.
20.2 PLANT TYPES 1 HABIT AND HABITAT

1. Annual A plant that completes its entire life cycle, from seed to reproduction to death, in
one single year.
2. Aquatic Organisms growing in water.
3. Biennial A plant that completes its entire life cycle in two years.
4. Climber A weak-stemmed plant with roots in the ground and climbing the other plants,
etc. to support itself with the help of its tendrils or adventitious roots, etc. Usually climbers
twist around their support.
5. Creeper A plant which is unable to support itself, and spreads along the ground.
1
Arranged alphabetically.
550 Terms of Plant Description 177
6. Epiphyte A plant, growing on stems or branches of other plants, with no roots in the ground,
but not taking nutrients on which it is growing.
7. Habit Appearance of an organism, e.g. herb, shrub, etc.
8. Habitat The place in which an organism or a community is found.
9. Herb A small, usually annual plant, with no wood in its stems or roots.
10. Liana An annual or perennial, elongate plant, with weak stem, which is often climbing.
11. Parasite An organism which takes all its nutrients from the tissues of another organism,
usually with harmful effects.
12. Perennial A plant that grows and reproduces for many years.
13. Shrub A much branched, small, woody perennial plant with several branches from ground
level upwards.
14. Tree A tall, woody, perennial plant with a single trunk which usually bears branches.
15. Vine A creeper of family Vitaceae.
16. Xerophyte A plant that lives in dry habitats or in a desert.
20.3 ROOT
20.3.1 Root Types1
1. Adventitious Root Root arising from any part of the plant body other than the radicle. It
arises from an organ other than the root.
2. Tap Root The main, persistent, primary root of the plant which shows apical dominance
and develops from the radicle.
20.3.2 Modified Roots1 (Fig. 20.1)

1. Aerial A root arising from a part of a plant which is above the ground, e.g. betel.
2. Annulated Root having a series of ring-like swellings, e.g. Psychotria.
3. Assimilatory Green, chlorophyll-containing roots, e.g. Trapa natans.
4. Buttress Roots having plank-like or board-like growth on the upper side. These are supportive
structures, e.g. Terminalia bellirica.
5. Clasping or Climbing These arise on the nodes of weak-stemmed plants, and help them to
climb or fix on the support, e.g. Pothos.
6. Conical Cone like, i.e. broad at the base and tapers gradually towards apex, e.g. Daucus
carota.
7. Contractile Roots capable of shortening and containing a wrinkled surface, e.g. Crocus.
8. Fasciculated Fleshy or tuberous roots in a cluster, e.g. Asparagus.
9. Fibrous Fine, thread-like roots, e.g. monocotyledons.
10. Fleshy Succulent roots, e.g. Dahlia.
1
178 Plant Taxonomy
Climbing Root
Annulated Root Assimilatory Root
Conical Root Contractile Root

Fasciculated Root Fusiform Root
Leaf Root
Moniliform Root Mycorrhizal Root
Napiform Root
Pneumatophores
Prop Root Tubercular Root
Fig. 20.1 Root modifications.
11. Fusiform Root thickened in the middle and tapering at the ends, e.g. Raphanus sativus.
12. Haustorial These are absorbing roots, present within the host of some parasitic plants, e.g.
Balanophora.
13. Knee or Aerating These are vertical or horizontal, aboveground roots, e.g. Heritiera.
14. Leaf Roots Roots arising on the margin of leaf from adventitious buds, e.g. Bryophyllum.
15. Moniliform Root thickened at certain intervals giving a beaded appearance, e.g.
Momordica.
16. Mycorrhizal Root in symbiotic relationship with fungal hyphae, e.g. Monotropa.
17. Napiform Root almost spherical at one end and tapering sharply at the lower part, e.g.
turnip, Beta vulgaris.
18. Nodulose Root bearing a small knot at or near the apex, e.g. Curcuma domestica.
19. Pneumatophorous or Respiratory Spongy, aerating roots of marshy plants, e.g.
Rhizophora.
20. Prop or Stilt Supportive roots which grow out from the bottom of the trunk into the ground,
e.g. Ficus benghalensis.
21. Tuberous or Tubercular Fleshy roots appearing like stem tubers, e.g. Ipomoea batatas
(Sweet Potato), Mirabilis jalapa.
20.4 STEM (Types, Surface, Forms and Modifications; Fig. 20.2)1

1. Acaulescent Stemless or with inconspicuous stem.
2. Aerial Stem which remains above the ground, e.g. Sesbania.
3. Angular Stem showing many angles in a transverse section, e.g. Asparagus.
4. Arborescent Woody and treelike, e.g. Mangifera.
5. Ascending Inclined upward.
6. Branched Stem with many branches, e.g. Ranunculus.
7. Bud Small embryonic stem tip bearing leaves or flowers, or both.
8. Bulb An underground, short, erect stem covered by fleshy leaves, e.g. Allium cepa.
9. Bulbel A young small bulb developed from the base of a large bulb.
10. Bulbil A bulblike body developed on the aboveground parts, e.g. Agave.
11. Caudex A short, hard, overwintering base of a perennial herb, e.g. palms.
12. Caulescent Possessing a distinct stem.
13. Cespitose Short, much branched, cushion-like plant.
14. Cladode A phylloclade in which a branch of a single internode becomes flat or leaf-like,
e.g. Asparagus, Ruscus.
15. Cladophyll A leaflike, flattened, green stem.
16. Climbing Growing towards upper side by means of tendrils, petioles, adventitious roots,
etc., e.g. pea, betel.
17. Columnar Erect with a strong trunk.
18. Corm Solid, round, bulblike, fleshy, underground stem, usually surrounded by membranous
scales, e.g. Gladiolus.
19. Cormel Young small corm developing at the base of parent corm, e.g. Amorphophallus.
20. Creeping Trailing stem having roots throughout its length, e.g. Oxalis.
21. Culm Flowering and fruiting stem of sedges (Cyperaceae) and grasses (Gramineae), e.g.
Cyperus, Cynodon.
1
180 Plant Taxonomy
Cormel
Bulbil
Bulb
Cladode
Corm
Offset
Decumbent Diffuse
Dichotomous
Procumbent
Repent
Phylloclade
Rhizome
Stolon Sucker
Supine
Runner
tendril
Tuber
Tendril Tiller Twinner
Fig. 20.2 Stem types and modifications

22. Cylindrical Stem showing circular outline in transverse section.

23. Decumbent Stem bending in one direction, e.g. Tridax.
24. Dichotomous Stem dividing equally into two, e.g. Pandanus.
25. Diffuse Branches of stem spreading in all directions, e.g. Boerhaavia.
26. Eramous With unbranched stems.
27. Erect Growing upright, e.g. Abutilon.
28. Fastigiate Strictly erect and parallel.
29. Fistular Stem hollow from inner side, e.g. wheat, bamboo.
30. Fruticose Woody and shrublike.
31. Geniculate Zigzag stem, or bending abruptly at node, e.g. Vitis.
32. Glabrous Smooth, or without hairs, e.g. Lemon.
33. Glaucous Shining and smooth.
34. Hairy Covered with hairs, e.g. Calotropis.
35. Herbaceous Soft, non-woody, dying to the ground at the end of the growing season, e.g.
Eclipta.
36. Internode Region of stem between any two nodes, e.g. Cynodon.
37. Laticiferous Containing latex, e.g. Calotropis.
38. Leaf Scar A mark on the stem showing former place of attachment of leaf base or
petiole.
39. Lenticel Lens-shaped or wartlike pore in the bark.
40. Node The area of the stem from which arise branches, leaves, or a leaf, e.g. sugarcane.
41. Offset A short, thick, creeping stem bearing a tuft of leaves above and a cluster of leaves
below at the apex, e.g. Eichhornia, Pistia.
42. Pachycauly Thick, short, frequently succulent stems, e.g. cacti.
43. Phylloclade A flat or round, green, succulent stem with leaves either ill-developed or modi-
fied into spines, e.g. Opuntia, Coccoloba.
44. Phyllode A flat petiole which has the appearance of a leaf, e.g. Australian Acacia.
45. Prickle A sharp, pointed extension of epidermis or cortex, e.g. Pisonia.
46. Primocane The first-year, nonflowering stem.
47. Procumbent or Prostrate Growing flat or parallel on the ground, e.g. Portulaca.
48. Pterocaulous Winged stem.
49. Pubescent Covered with fine silky hairs.
50. Ramose Branched stem.
51. Repent Creeping stem.
52. Rhizome A horizontal, prostrate, or underground stem bearing scale-like leaves, e.g. ginger,
Canna.
53. Rootstock A caudex, or a rhizome, or any underground portion of a plant, e.g. Alocasia.
182 Plant Taxonomy
54. Runner A horizontal stem, creeping aboveground, usually rooting and producing plants at
the nodes, e.g. Cynodon.
55. Sarcocauly Fleshy stems.
56. Scandent Climbing stem.
57. Scape A leafless flowering stem arising from an underground stem, e.g. Canna.
58. Scapose Possessing a scape.
59. Scars Remains of a point of attachment of leaf, stipule, scale, bud, etc.
60. Sclerocauly Hard, woody stems.
61. Spine A stiff, pointed outgrowth.
62. Stolon A runner, or a horizontal stem rooting at the nodes, e.g. Fragaria indica.
63. Stoloniferous Bearing sto1ons.
64. Subterranean Underground.
65. Sucker A subterranean creeping stem, usually fast-growing and adventitious, e.g. Mentha,
banana.
66. Suffrutescent Woody at the base.
67. Supine Prostrate stem with parts oriented upward.
68. Tendril Long, twisting appendage adapted for climbing, e.g. Vitis.
69. Thorn A sharp, pointed, reduced branch, e.g. Duranta.
70. Tiller A grass shoot developed from the base of the stem.
71. Trailing Stem sprawling on the ground with the help of adventitious roots.
72. Tuber Thick, enlarged fleshy tip of an underground stem, e.g. Solanum tuberosum.
73. Turion An overwintering bud.
74. Twig A short lateral branch of a woody stem.
75. Twinner Stem ascending by coiling on the support without any special device, e.g. Abrus.
76. Virgate Long, straight, slender or thin, stick-like.
77. Woody Hard in texture and possessing secondary xylem, e.g. Mangifera indica.
20.5 LEAF
20.5.1 Apex of Leaf (Fig. 20.3 A–M)
1. Acuminate Drawn out in the form of a long slender tail, or tapers to a protracted point
(Fig. 20.3 A).
2. Acute Ending into a sharp point in the form of an acute angle but not drawn out
(Fig. 20.3 B).
3. Apiculate Ending into a short, sharp, flexible point, or an apicula (Fig. 20.3 C).
4. Aristate Tapering to a very narrow, much elongated apex, or bearing a stiff awn
(Fig. 20.3 D).
5. Caudate Containing a tail-like appendage (Fig. 20.3 E).
F D E A G J
L
H
I C
B
M K
O
T
Y P
N R
U Q X V W
Fig. 20.3 A-M, Leaf apices; N-Y, Leaf bases.
6. Cirrhose Apex ending into a tendril like structure (Fig. 20.3 F).
7. Cuspidate Abruptly and sharply concavely constricted into a sharp and elongated pointed
tip (Fig. 20.3 G).
8. Emarginate Containing a deep or shallow notch at the apex (Fig. 20.3 H)
9. Mucronate Terminating abruptly into a short and sharp point (Fig. 20.3 I).
10. Mucronulate Diminutive or smaller form of mucronate (Fig. 20.3 J).
11. Obcordate Deeply lobed at the apex (Fig. 20.3 K).
12. Obtuse or Rounded Blunt or rounded (Fig. 20.3 L).
13. Retuse Slightly notched obtuse apex (Fig. 20.3 M).
20.5.2 Base of Leaf 1 (Fig. 20.3 N–Y)

1. Attenuate Base showing a long gradual taper (Fig. 20.3 N).
2. Auriculate Bearing an appendage or ear-shaped part at the base (Fig. 20.3 O).
3. Connate-perfoliate Base of opposite sessile leaves, the stem appears passing through them
(Fig. 20.3 P).
4. Cordate Heart-shaped (Fig. 20.3 Q).
5. Cuneate Wedge-shaped; triangular with the narrow end at the attaching point
(Fig. 20.3 R).
6. Hastate Resembling with an arrowhead; halberd-shaped (Fig. 20.3 S).
7. Oblique Two sides of base being unequal-sized (Fig. 20.3 T).
8. Obtuse Rounded or blunt (Fig. 20.3 U).
9. Peltate Attached to its stalk inside the margin (Fig. 20.3 V).
10. Perfoliate Sessile leaf whose base completely surrounds the stem (Fig. 20.3 W).
1
184 Plant Taxonomy
11. Sagittate Base like an arrowhead, or triangular (Fig. 20.3 X).

12. Truncate Blunt, or appearing as if cut off at the end (Fig. 20.3 Y).
20.5.3 Kinds, Modifications and Parts of Leaf 1

1. Blade Expanded, flat part of leaf.
2. Bract Reduced and modified leaf found in inflorescence.
3. Bracteole Small leaf found on the pedicel of flower.
4. Cauline Leaf arising only on the main axis.
5. Cauline and Ramal Leaves arising on the main axis as well as on side branches.
6. Chaff Bract present at the base of tubular flower, as in Asteraceae.
7. Cotyledon Embryonic leaf.
8. Epetiolate Sessile leaves.
9. Epetiolulate Sessile leaflets.
10. Epicalyx Leaves present below the true calyx, e.g. Malvaceae.
11. Evergreen Persistent.
12. Exstipulate Without stipules.
13. Floral leaves Calyx, corolla, stamens and carpels.
14. Fly trap Hinged leaves of insectivorous plants.
15. Fugacious Falling soon.
16. Glume Bract of the spikelets of sedges and grasses.
17. Incomplete Leaf without one or more parts, e.g. without petiole.
18. Involucral bracts Bracts present below the inflorescence in Asteraceae.
19. Leaflet A distinct or separate segment of leaf.
20. Lemma Outer bract subtending the grass floret.
21. Ligule Finger-like small part present at the upper end of leaf sheath.
22. Pale Chaff.
23. Palea Inner bract subtending the grass floret.
24. Parts of leaf Leaf base, stipule, petiole, lamina, midrib, veins, veinlets.
25. Petiolate Leaf with a distinct stalk or petiole.
26. Petiolulate Leaflets with a stalk.
27. Phyllary A bract subtending the inflorescence in Asteraceae.
28. Pitcher Tubular or pitcher-shaped leaf of insectivorous plants.
29. Prophyll Bracteole.
30. Pulvinus Swollen base of petiole or petiolule.
31. Rachilla Secondary axis of a compound leaf.
1
32. Rachis Main axis of a pinnately compound leaf.

33. Radical Leaves arising from bulb or other underground stem.
34. Scaly leaf Small, nongreen leaf.
35. Seed leaves Cotyledons.
36. Sessile Without petiole.
37. Spathe An enlarged bract enclosing an inflorescence.
38. Sporophylls Spore-bearing leaves.
39. Stipel Scale or gland present at the base of petiolule.
40. Stipule Scale, gland or blade-like structure at the base of petiole of leaf.
41. Storage leaf Fleshy, succulent leaf.
42. Tendril Coiled, modified leaflet.
20.5.4 Margin of Leaf 1 (Fig. 20.4)

1. Biserrate When each tooth of the serrated margin is again serrated.
2. Ciliate Margin bearing hairs.
3. Crenate With blunt, low rounded teeth.
4. Crenulate Finely crenate.
5. Crispate Curled and extremely undulate.
6. Dentate With sharp, marginal teeth pointing outward.
7. Dissected Cut or deeply divided into many segments.
8. Divided Cut into distinct segments or sections, extending to the midrib or base of the
leaf.
9. Entire With a continuous smooth margin; lacking any teeth, lobes or indentations.
10. Incised Provided with sharp and irregular incisions.
11. Lacerate Torn or irregularly cut.
12. Lobed Provided with many lobes extending one third to one-half the distance between the
midrib and the margin.
13. Palmatifid Cut about halfway down in a palmate form.
14. Parted Dissected or cut almost to the midrib.
15. Pinnatifid Parted in a pinnate way, or divided almost to the midrib.
16. Repand Wavy in appearance.
17. Revolute With the margin rolling inward i.e. towards the underside of the leaf.
18. Runcinate When the teeth of the serrated margin are pointing backward.
19. Serrate With the marginal teeth pointing towards the apex.
20. Serrulate When the teeth in the serrated margin are very minute.
1
186 Plant Taxonomy
Biserrate Ciliate Crenate Crispate Dentate Entire Incised Lacerate Lobed
Parted Revolute
Palmatifid Pinnatifid Serrate Undulate
Serrulate
Fig. 20.4 Leaf margins.
21. Sinuate With a deeply wavy margin.

22. Spinous Provided with projecting spines.
23. Undulate With a slightly wavy margin.
20.5.5 Phyllotaxy 1
1. Alternate Bearing one leaf at each node.
2. Distichous When alternate leaves appear on just two sides of the stem.
3. Opposite Bearing leaves paired at each node on opposite sides.
4. Opposite decussate When two successive opposite pairs of leaves occur at right angle to
each other.
5. Opposite superposed When all the successive opposite pairs of leaves occur at the same
plane.
6. Radical or basal When leaves often form a cluster at the ground level.
7. Tristichous, pentastichous and octostichous In the alternate arrangement of leaves, if the
fourth leaf comes over the first one, the arrangement is called tristichous; if the sixth leaf
comes over the first one after completing two revolutions of spiral, it is called pentastichous;
and if the ninth leaf comes over the first one after completing three revolutions of the spiral,
it is called octostichous.
8. Whorled or verticillate Bearing three or more leaves at each node.
1
20.5.6 Shape 1 of Leaf (Fig. 20.5)

1. Acicular Needle shaped.
2. Auriculate Ear shaped.
3. Cordate Heart shaped.
4. Cuneate Wedge-shaped with lower end narrow.
5. Deltoid Delta-like or triangular.
6. Elliptical Oval in outline, being narrowed to form rounded ends and widest at the
middle.
7. Falcate Sickle-shaped.
8. Filiform Long, slender and thread-like.
9. Hastate Like an arrowhead.
10. Lanceolate Much longer than broad; or lance-shaped; or widening above the base and
tapering towards the tip.
11. Linear Long, flat, and narrow with almost parallel sides.
12. Lorate Strap-shaped, or like a narrow strip of leather.
13. Lyrate Like a lyre, i.e. having a big terminal lobe and several smaller lateral lobes.
14. Obcordate Inversely cordate.
15. Oblanceolate Broad at the middle and tapering towards both the ends.
16. Oblique When two halves of the lamina are unequal.
17. Oblong Long, wide, with the parallel margins.
18. Obovate Upper terminal half broader than the lower basal half; opposite to ovate.
19. Ovate Egg shaped; broad at the base and narrowing towards apex.
20. Pandurate Resembling obovate with a distinct concavity along each basal side; fiddle-
shaped.
21. Pedate Like the claw of a bird; or palmately lobed.
22. Reniform Kidney-shaped.
23. Rhomboid Rhombic-shaped.
24. Rotund Nearly circular.
25. Runcinate Irregularly serrate or sharply incised with the teeth pointing towards the base.
26. Sagittate Like an arrowhead; or triangular.
27. Spathulate Spoon-shaped.
28. Subulate Tapering from base to apex; awl-shaped.
20.5.7 Types of Stipule (Fig. 20.6)

1. Adnate Attached fully to petiole as wings.
2. Basal Stipules attached near the base of the petiole.
1
188 Plant Taxonomy
Acicular Auriculate Cordate Cuneate Elliptical Falcate
Hastate Lanceolate Linear Lorate Lyrate Obcordate
Oblanceolate Oblique Oblong Obovate Ovate Pedate
Reniform Rotund Sagittate Spathulate Subulate
Fig. 20.5 Leaf shapes.
3. Foliaceous Stipule resembling with a large leaf.

4. Interpetiolar Present in between the petioles of opposite leaves.
5. Intrapetiolar Situated in the axil of leaves, and their margins fuse above the petiole.
6. Lateral Present laterally on the petiole.
adnate tendrillar
intrapetiolar
ventral
lateral
foliaceous
interpetiolar
ochreate
Fig. 20.6 Types of stipule.
7. Ochreate Fused to form an ochrea or a fused tubular nodal sheath around the internode.
8. Photosynthetic Large, green and leaflike or foliaceous.
9. Scaly Dry, small and membranous stipule.
10. Sheathing or Protective Enclosing a bud or flower like a sheath.
11. Spinous Stipule in the form of a hard spine.
12. Tendrillar Stipule modifying into a tendril.
13. Ventral Present on the ventral side of the petiole.
14. Vestigial Very small, minute, or remnant.
20.5.8 Types of Leaf (Fig. 20.7)

1. Compound Leaf A leaf, in which the leaf blade or lamina remains divided into smaller,
bladelike parts or leaflets is called a compound leaf. A compound leaf may be palmately
compound or pinnately compound.
2. Palmately Compound Leaf If the leaflets diverge from a common point at the end of the
petiole, in the same way as the fingers from the palm of the hand, the leaf is called palmately
compound. In such leaves, if a single leaflet is articulated to the petiole, it is called unifoliate;
190 Plant Taxonomy
leaflet
bud
Palmately Compound Leaf Unifoliate Bifoliate Trifoliate
bud
Unipinnate Unipinnate
Quadrifoliate Multifoliate (paripinnate) (imparipinnate) Bipinnate
bud bud
Tripinnate Decompound Simple leaf
Fig. 20.7 Types of leaf.
if two, three or four leaflets are articulated to the petiole, it is called bifoliate, trifoliate or
quadrifoliate, respectively; and if five or more leaflets are articulated to the petiole, it is
called multifoliate.
3. Pinnately Compound Leaf If the leaflets are attached on both sides of one central rachis,
the leaf is called pinnately compound. In such leaves, if the leaflets are attached directly on
the midrib, the leaf is called unipinnate. A unipinnate leaf having even number of paired
leaflets is called paripinnate, while that which contains an odd terminal leaflet is called
imparipinnate. Such a pinnately compound leaf, in which the midrib produces secondary
axis, and on the latter are present the leaflets, is called bipinnate; if the midrib of pinnately
compound leaf produces secondary axis, and the latter produces the tertiary axis which
bears the leaflets, it is called tripinnate. If the leaf is more than thrice pinnate, it is called
decompound.
4. Simple Leaf A leaf with the blade in a single part is called simple leaf. The single blade
may, however, be variously divided.
20.5.9 Venation of Leaf (Fig. 20.8)

1. Venation The pattern of veins on the surface of a leaf.
2. Parallel When the veins run parallel to each other in the lamina of the leaf, the venation is
called parallel, e.g. monocots. In this type of venation, if only one principal vein is present,
it is called unicostate, and if several principal veins are present, it is called multicostate.
3. Reticulate When the pattern of the veins in the lamina of the leaf is like a network, e.g.
dicots.
4. Unicostate and Multicostate In both parallel and reticulate venations, if only one principal
vein is present, it is called unicostate, and if several principal veins are present, it is called
multicostate. In the multicostate types, if the veins run in a curved manner from the base
of the blade to its apex, it is called convergent, and if the veins arise at the base of the leaf
blade and then diverge from one another towards the leaf margin, it is called divergent.
A B
D E F
Fig. 20.8 Leaf venation. A: Unicostate parallel; B: Unicostate reticulate; C: Multicostate parallel
convergent; D: Multicostate parallel divergent; E: Multicostate reticulate convergent; F: Multicostate
reticulate divergent.
192 Plant Taxonomy
20.6 INFLORESCENCE
20.6.1 Basic Categories
Cymose, racemose, and special types are three basic categories of inflorescences (mode of the
arrangement of the flowers on the floral shoot) of angiosperms.
1. Cymose In this type the growth of the main axis is checked soon by the development of a
flower at the apex, and the lateral axis below the terminal flower also ends in a flower, and
thus its growth is also checked. In cymose inflorescence, the terminal flower is the oldest
and the young flowers are present on the lower side. Helicoid, circinnus, rhipidium, dichasium
and polychasium are some of the examples of cymose.
2. Racemose In this type, the main axis does not terminate into a flower, but it keeps on
growing continually and gives off flowers laterally in acropetal succession. Here the youngest
flower is present at the apex and the older flowers towards the base. Raceme, spike, spikelet,
panicle, catkin, spadix, corymb, umbel and capitulum or head are some of the examples of
racemose.
3. Special Types Cyathium, verticillaster, and hypanthodium are some special types of
inflorescences.
20.6.2 Types of Inflorescence 1 (Fig. 20.9)

1. Catkin A spike-like, deciduous, elongate, inflorescence, with scaly bracts, and unisexual,
apetalous and sessile flowers.
2. Cincinnus A modified helicoid cyme having the short pedicels on the developed side.
3. Compound Corymb A branched corymb having pedicellate flowers.
4. Compound Cyme A branched cyme having pedicellate flowers.
5. Compound Umbel A branched umbel having primary rays arising at a common point with
a secondary umbel arising from the tip of the primary rays.
6. Corymb A raceme whose lower stalks are longer than the upper ones, so the inflorescence
has a flat top.
7. Cyathium A cup-shaped involucre containing nectar-secreting glands, a centrally-placed
large female flower, and many male flowers.
8. Dichasium or Simple Cyme Determinate, dichotomous inflorescence of pedicellate flowers
having pedicels of equal length.
9. Glomerule An indeterminate inflorescence having dense cluster of sessile or subsessile
flowers.
10. Head or Capitulum A dense cluster of several sessile or subsessile flowers on a compound
receptacle or torus.
11. Helicoid Cyme Curved and unbranched inflorescence of pedicellate flowers having the
branches only on one side.
1
Terms arranged alphabetically.
Cincinnus
Scorpioid Simple
Helicoid Compound Simple
Compound Compound
CYMES CORYMBS DICHASIA
HEADS UMBELS
Indeterminate
Determinate Determinate Indeterminate Compound
PANICLE RACEME SPIKE
CATKIN GLOMERULE HYPANTHODIUM

SCAPE SECUND
spathe
SPIKELET
SPADIX THYRSE UMBEL
male flower
involucre
nectary
female flower
VERTICILLASTER
CYATHIUM
Fig. 20.9 Types of inflorescence.

194 Plant Taxonomy
12. Hypanthodium An inflorescence having sessile flowers on the wall of a concave capitulum,
opening by a small ostiole. Male flowers are situated near the periphery and female flowers
in the centre, e.g. Ficus.
13. Panicle Branched inflorescence with pedicellate flowers arranged in the form of number
of racemes.
14. Pleiochasium It is a compound dichasium in which each dichasium has three lateral
branches. It is also called polychasium or multiparous cyme.
15. Raceme An indeterminate, unbranched inflorescence with a single axis and the flowers
arranged along the main axis on pedicels.
16. Scorpioid Cyme A zig-zag cymose inflorescence that appears to coil like a scorpion’s tail.
In this type, the branches develop alternately on opposite sides of the rachis.
17. Secund Flowers arranged only on one side of the rachis.
18. Solitary Axillary Single-flowered; flower attached in the axis.
19. Solitary Terminal Single-flowered; flower attached at the apex, and not in the axis.
20. Spadix A fleshy or thick, spikelike inflorescence with very small flowers usually enclosed
in a spathe.
21. Spike Elongate, unbranched, indeterminate inflorescence with sessile flowers.
22. Spikelet A small spike, the basic inflorescence unit of Cyperaceae and Gramineae.
23. Thyrse A compact and compound panicle having an indeterminate main axis and laterally
determinate axes.
24. Umbel An inflorescence in which all the pedicels are of the same length and arise from
the same point.
25. Umbellet A secondary umbel in a compound umbel.
26. Verticillaster Whorled dichasial cymes arranged at the nodes of an elongate axis, e.g.
Labiatae.
20.7 FLOWER PARTS AND TYPES 1

1. Accessory Whorls Calyx and corolla.
2. Achlamydous Without calyx and corolla.
3. Actinomorphic Flowers with radial symmetry; or flowers which can be bisected into similar
halves along two or more planes.
4. Amorphic Flowers without symmetry.
5. Androecium All stamens of a flower.
6. Androgynophore Stripe bearing both androphore and gynophore together.
7. Androphore Elongated internodal part between corolla and androecium.
8. Anthophore Elongated internodal part between calyx and corolla.
9. Bisexual or Hermaphrodite Flowers having both the sex organs.
1
Terms arranged alphabetically,
10. Bract Modified leaf which develops in the axil of the flower.
11. Bracteate With bract.
12. Bracteolate With bracteole.
13. Bracteole Small leaf borne on pedicel of flowers.
14. Calyx Whorl of sepals.
15. Carpel A unit of gynoecium; or floral organ that bears ovules.
16. Complete Flowers having all the four floral whorls, i.e. sepals, petals, androecium and
gynoecium.
17. Corolla Whorl of petals.
18. Dimerous, Trimerous, Tetramerous and Pentamerous Flowers in which various floral parts
are arranged in groups of two, three, four and five, are called dimerous, trimerous, tetramer-
ous and pentamerous, respectively.
19. Ebracteate Without bracts.
20. Epicalyx Leaves resembling sepals below the true calyx.
21. Epigynous Flowers with inferior ovary; or those having the floral parts situated above the
ovary.
22. Essential Whorls Androecium and gynoecium.
23. Gynoecium or Pistil Group of all carpels of a flower.
24. Gynophore Elongated axis between androecium and gynoecium.
25. Haplomorphic Petals or tepals coloured.
26. Hypanthium Fused basal portion of sepals, petals or stamens around the ovary.
27. Hypogynous Flowers with superior ovary; or those having the floral parts situated below
the ovary.
28. Monochlamydous Flowers with only one whorl.
29. Monoecious Having separate male and female flowers on the same individual.
30. Pedicel The stalk of the flower.
31. Pedicellate Flower with pedicel.
32. Perianth When there is no differentiation of calyx and corolla.
33. Perigynous Flowers with half-inferior ovary; or those having floral parts situated around
the ovary.
34. Petal Individual unit of corolla.
35. Pleomorphic Actinomorphic flowers with number of reduced parts.
36. Polyphore A receptacle having several distinct carpels.
37. Receptacle, Thalamus or Torus Tip of the axis bearing floral appendages.
38. Sepal Individual unit of calyx.
39. Sessile Without stalk.
40. Stamen Individual unit of androecium.
196 Plant Taxonomy
41. Tepal Individual unit of perianth.

42. Unisexual Flower with only one sex, i.e. either male or female.
43. Zygomorphic Flowers with bilateral symmetry; or flowers which can be bisected into similar
halves in only one plane; or flowers with parts usually irregular and reduced in number.
20.8 CALYX 1
1. Anterior Lobes Abaxial lobes; or lobes that remain away from axis.
2. Aposepalous With separate sepals.
3. Asepalous Without sepals.
4. Bilabiate Consisting of two lips.
5. Caducous Sepals which wither or fall soon.
6. Campanulate Bell-shaped.
7. Chorisepalous With separate sepals.
8. Cupulate Cup-like.
9. Dorsal Side Abaxial, or back, or lower side of the sepals.
10. Fringed Modified margin of sepals or petals.
11. Gamosepalous Fused sepals.
12. Infundibuliform Funnel shaped.
13. Pappus Reduced, scaly or hairy calyx of Asteraceae.
14. Persistent Sepals which persist even in the fruit.
15. Petalloid Coloured sepals, except green.
16. Polysepalous When sepals are free.
17. Posterior Lobe Adaxial lobe; or lobe that remains next to axis.
18. Sepal A unit of calyx.
19. Spurred When one or more sepals are produced into spur.
20. Synsepalous With fused sepals.
21. Tubular Like a tube; cylindrical.
22. Urceolate Urn-shaped.
23. Ventral Side Upper or top side of sepal or petal.
20.9 AESTIVATION 1 Fig. 20.10)

1. Aestivation The way in which sepals and petals are arranged in the bud condition.
2. Imbricate Aestivation Out of the five sepals or petals, one external, one internal, and other
three are partly external and partly internal.
1
Imbricate Quincuncial Twisted Valvate Vexillary
Fig. 20.10 Types of aestivation.
3. Quincuncial Aestivation Out of the five sepals or petals, two external, two internal, and
remaining one is partly external and partly internal.
4. Twisted Aestivation When margins of each part are overlapped regularly, i.e. one edge of
the sepal or petal are overlapped by the preceding part.
5. Valvate Aestivation When the sepals or petals meet edge to edge without overlapping each
other.
6. Vexillary Aestivation Out of the five sepals or petals the posterior one is the largest and
covers the two lateral sepals or petals, and the latter in turn overlap the two smallest and
anterior sepals or petals, e.g. Papilionaceae.
20.10 COROLLA
20.10.1 General Terms 1
1. Apetalous Without petals.
2. Apopetalous Having separate petals.
3. Carina Keel, or two fused petals of a papilionaceous flower.
4. Choripetalous Apopetalous.
5. Gamopetalous With fused petals.
6. Lodicule Scale-like perianth part of Gramineae.
7. Petal An individual unit of corolla.
8. Polypetalous With free petals.
1
198 Plant Taxonomy
9. Standard or Vexillum Wider petal in a papilionaceous flower.

10. Sympetalous With fused petals.
11. Wing Lateral petals, e.g. Fabaceae.
20.10.2 Types of Corolla 1 (Fig. 20.11)

1. Bilabiate or Bilipped Zygomorphic gamopetalous corolla with two unequal lips or
divisions.
2. Calcarate Spurred corolla.
3. Campanulate Bell-shaped.
4. Carinate Keeled.
Bilabiate
Campanulate Cruciform
Caryophyllaceous
Hypocrateriform
Infundibuliform Ligulate Papilionaceous
Personate
Rosaceous Rotate
Tubular Urceolate
Fig. 20.11 Types of corolla.
1
5. Caryophyllaceous When all the five petals of the corolla are clawed and have their limbs
spreading out.
6. Cruciate or Cruciform All the four petals are arranged in the form of a cross, e.g.
Cruciferae.
7. Hypocrateriform Gamopetalous, salver-shaped corolla i.e. basal portion narrow and tubular
having abruptly expanding flat apical portion.
8. Infundibuliform Funnel-shaped.
9. Ligulate Zygomorphic, gamopetalous, strap-shaped corolla with a short narrow tube and
strap-like upper portion.
10. Papilionaceous or Butterfly-like When five petals of the corolla are arranged in a butterfly
like shape. It consists of a posterior largest vexillum, two lateral alae, and two posterior fused
petals called keel, e.g. Papilionaceae.
11. Personate It is also a bilabiate corolla but here the corolla mouth is closed because two
lips are very close to one another.
12. Rosaceous When five or more petals are spreading like those of rose, e.g. Rosaceae.
13. Rotate Wheel-shaped gamopetalous corolla having narrow corolla tube, and the limbs of
petals are at right angle to the lobe.
14. Salverform Trumpet-shaped.
15. Tubular Cylindrical; petals fused to form a tube.
16. Urceolate Urn- or pitcher-shaped.
20.11 PERIANTH 1
1. Gamotepalous With fused tepals.
2. Lodicule Scalelike perianth part of Gramineae.
3. Perianth Collective term applied for calyx and corolla.
4. Petalloid Resembling with petals.
5. Polytepalous With free tepals.
6. Sepalloid Resembling with sepals.
7. Tepal Individual unit of perianth.
20.12 ANDROECIUM 1 Fig. 20.12

1. Adnate When filament runs throughout the entire length of the anther from the base to the
top (Fig. 20.12).
2. Allagostemonous Condition in which stamens are attached to petals and torus alternately.
3. Alternipetalous Stamens present alternately with the petals.
4. Anther Pollen-bearing portion of stamen.
1
200 Plant Taxonomy
anther
stamens
connective
stamen
filament
A Stamen Diadelphous Didynamous Gynandrous

A B C D
stamen
Polyadelphous Synandrous Syngenesious

Monadelphous
E F G H
stamen
Tetradynamous Adnate Basifixed Dorsifixed Versatile

I J K L M
Fig. 20.12 A–I: Types of androecium; J–M: Fixation of filament.
5. Antipetalous Stamens opposite to petals.

6. Antiphyllous Stamens opposite to tepals.
7. Antisepalous Stamens opposite to sepals.
8. Apostemonous With separate stamens.
9. Appendicular Stamen with a modified or protruding connective.
10. Basifixed Filament attached with the base of the anther lobe.
11. Connective Filament extension of the anther between thecae.
12. Diadelphous With the filaments of stamens united or connate in two groups with their
anthers being free.
13. Diandrous Flowers with two stamens.

14. Didymous With stamens in two equal pairs.
15. Didynamous When two out of the four stamens are larger and remaining two are smaller
in size (Fig. 20.12).
16. Diplostemonous When stamens are arranged in two whorls, of which the outer whorl is
opposite to sepals and inner whorl is opposite to petals.
17. Dorsifixed When the filament is attached on the dorsal side of the anther lobes
(Fig. 20.12).
18. Epipetalous When the stamens remain attached to the petals.
19. Epiphyllous When the stamens remain attached with the tepals.
20. Episepalous When the stamens remain attached with the sepals.
21. Extrorse Stamen which dehisces longitudinally outward.
22. Filament Stalk of the stamen.
23. Gynandrous When stamens are fused with gynoecium.
24. Introrse Stamen which dehisces longitudinally inward.
25. Laminar When stamens are leaflike with no distinct anther and filament.
26. Laterose Stamen which dehisces longitudinally and laterally.
27. Locule Chamber of an anther.
28. Monadelphous When filaments of one group of stamens are fused but their anthers are
free (Fig. 20.12).
29. Monandrous Flower with one stamen.
30. Obdiplostemonous When the stamens are arranged in two whorls, of which the outer whorl
of stamens being opposite to petals.
31. Petalloid Stamens resembling with petals, showing no distinction between anther and
filament.
32. Petalostemonous With filaments of stamens fused to petals with free anthers.
33. Phaenantherous With exerted stamens.
34. Pollen grain Young male gametophyte.
35. Pollen sac Male sporangium.
36. Polyadephous When filaments are united in several groups with their anthers remaining
free (Fig. 20.12).
37. Polyandrous With several free stamens.
38. Poricidal Dehiscing through a pore at the theca apex.
39. Stamen Pollen-bearing floral organ of angiosperms.
40. Staminode Sterile stamen with reduced anthers.
41. Synandrous With anthers as well as their filaments being united in the form of one
group.
202 Plant Taxonomy
42. Syngenesious With fused anthers and free filaments (Fig. 20.12).
43. Tetrandrous Flowers with four stamens.
44. Tetradynamous Two out of a total of six stamens are short while the remaining four are
long.
45. Theca Half of the portion of anther containing two pollen sacs.
46. Triandrous Flowers with three stamens.
47. Tridynamous Stamens arranged in two equal groups of three.
48. Valvular When the stamen dehisces through a pore covered by a flap of tissue.
49. Versatile Dorsifixed but it appears as if the anther is swinging freely on the filament
(Fig. 20.12).
20.13 GYNOECIUM 1 Fig. 20.13
20.13.1 Parts, Fusion Types, Carpel Parts and Types, Style Types,
and Ovary Position
1. Apocarpous With free or separate carpels.
2. Astylocarpellous Without a stipe and a style.
3. Astylocarpepodic With a stipe and without a style.
4. Astylous Without any distinct style.
5. Carpel Organ of the flower that bears ovules.
6. Epigynous The condition in which the sepals, petals and androecium are attached to the
floral tube above the ovary.
7. Funiculus The stalk with which the ovule is attached to the placentum.
8. Geniculate Style The style which bents abruptly.
9. Gynobasic Style The style which is attached at the base of the ovary in central
depression.
10. Heterostylous With styles of different shapes or lengths.
11. Homostylous With styles of same shapes or lengths.
12. Hypogynous The condition in which sepals, petals and androecium are attached to the floral
tube below the ovary.
13. Inferior Ovary The ovary in the epigynous condition.
14. Locule The cavity of the ovary.
15. Monocarpellary to Polycarpellary With one to several carpels; bicarpellary if two carpels
are present, tricarpellary if three carpels are present, and tetra-and pentacarpellary if four
and five carpels are present.
16. Ovary Ovule-bearing part of the gynoecium.
1
stigma
stigma style
stigma stigma
ovary style ovary
A ovary ovary
stipe stipe
pedicel pedicel pedicel
Astylocarpellous Astylocarpepodic Stylocarpellous Stylocarpepodic
Apocarpous Semicarpous Syncarpous Synovarious Synsty- Unicarpellous

lovarious
Geniculate Gynobasic Heterostylous Homostylous Terete Fimbriate

chalaza funiculus
antipodal cell
polar nuclei stigma
nucellus
outer style
locule
integument
placenta
inner embryo sac
integument egg stipe
synergid
micropyle funiculus E
D
Fig. 20.13 A: Types of carpel; B: Types of gynoecium; C: Types of style; D: V.s. of an ovule and;
E: V. s. of a carpel
17. Ovule Young seed made up of nucellus and integuments.

18. Parts of a Carpel Funiculus, locule, ovary, ovule, placenta, stigma, style.
19. Parts of Gynoecium Carpel, locule, ovary, ovule, placenta, stigma, style.
20. Parts of Ovule Chalaza, embryo sac, integuments, micropyle, nucellus, raphe.
21. Perigynous The condition in which the sepals, petals and androecium are attached to the
floral tube around the ovary.
22. Pistillode A sterile carpel.
23. Placenta Ovule-bearing region of the wall of ovary.
24. Raphe Part of an ovule present in the form of a longitudinal ridge on its outer
integument.
25. Replum False septum.
26. Semicarpous With fused ovaries of adjacent carpels and their free styles and stigmas.
204 Plant Taxonomy
27. Semi-inferior The position of ovary in the perigynous condition.

28. Stigma Uppermost, pollen-receptive part of the gynoecium.
29. Stipe Stalk of the gynoecium.
30. Style Long portion of gynoecium between ovary and stigma.
31. Stylocarpellous Carpel without a stipe and with a style.
32. Stylocarpepodic With a stipe and a style.
33. Stylodious With single free carpel.
34. Superior The position of ovary in the hypogynous condition.
35. Syncarpous With fused carpels.
36. Synovarious The condition when the ovaries of adjacent carpels are fused while their styles
and stigmas separate.
37. Synstylovarious The condition when the ovaries and styles of adjacent carpels are fused
while their stigmas separate.
38. Terete Cylindrical and elongate.
39. Unicarpellous With single carpel.
40. Unilocular to Multilocular Ovary with one chamber or locule is called unilocular; with
two chambers is called bilocular; with three and four chambers are called trilocular and
tetralocular; and with many locules is called multilocular (Fig. 20.14).
locules locule locule ovule
ovules
ovules
A B C D E
Fig. 20.14 Chambers of the ovary—A: Unilocular; B: Bilocular; C: Trilocular; D: Tetralocular, and;
E: Pentalocular.
20.13.2 Types of Stigma (Fig. 20.15)

1. Capitate Like a cap or head.
2. Clavate Club-shaped.
3. Crested Possessing a terminal tuft or ridge.
4. Decurrent Long and extending downward.
5. Diffuse Spread over a wide surface.
6. Discoid Like a disc.
7. Fimbriate Fringed.
8. Lineate In the form of small lines.
9. Lobed Divided into some lobes.
Capitate Clavate Crested Decurrent Diffuse
Discoid Fimbriate Lineate Lobed Plumose Terete
Fig. 20.15 Types of stigma.
10. Plumose Like a feather.

11. Terete Elongate and cylindrical.
20.14 PLACENTATION 1 Fig. 20.16

1. Axile The placentae develop along the central axis in a compound ovary with septa. It
occurs in a bilocular to multilocular ovary.
2. Basal The placenta develops at the base of the ovary, i.e. directly on the thalamus. It occurs
in a unilocular ovary.
3. Free-central The placenta develops along the central axis in a compound ovary without
septa. It occurs in unilocular ovary.
4. Laminar, or Laminate, or Superficial The placenta develops over the inner surface of the
ovary wall. It occurs in a multicarpellary ovary.
5. Marginal The placenta develops along the margin of the simple ovary. It occurs in mono-
carpellary and unilocular ovary.
6. Parietal The placentae develop on the wall or intruding portions of a compound ovary. It
occurs in bicarpellary to multicarpellary but unilocular ovary.
7. Pendulous or Apical The placenta develops at the top of the ovary and the ovules remain
suspended in a pendulous manner.
Axile Basal Free-central Laminate Marginal Parietal Pendulous
Fig. 20.16 Types of placentation.
1
206 Plant Taxonomy
20.15 FRUIT 1 Figs. 20.17, 20.18, 20.19

1. Achene Developing from a monocarpellary superior ovary, the achene is a one-seeded, dry,
indehiscent, simple fruit having its seed attached to fruit wall only at one point, e.g. Mirabilis,
Clematis.
2. Amphisarca A fleshy, simple, berry-like fruit with a woody rind, e.g. Lagenaria.
3. Balausta A simple, dry, many-seeded and many-loculed, indehiscent fruit with a tough
pericarp, e.g. Punica.
4. Berry Fleshy, simple fruit with succulent pericarp, e.g. tomato, banana, Vitis.
5. Bibacca A multiple type of fruit formed as a fused double berry, e.g. Lonicera.
6. Cacervulus A dry, schizocarpic fruit which develops from bi- to multicarpellary, syncar-
pous, superior ovary with many locules, e.g. Labiatae.
7. Calybium A hard, unichambered, dry, indehiscent fruit derived from an inferior ovary, e.g.
Quercus.
8. Capsule A dry, dehiscent fruit derived from a bi- to multicarpellary superior or inferior
ovary, e.g. Papaver.
9. Caryopsis A dry, indehiscent fruit derived from monocarpellary ovary in which pericarp
is inseparably fused with testa, e.g. Gramineae.
10. Cremocarp A dry, schizocarpic fruit derived from bicarpellary, syncarpous, inferior ovary
with one ovule in each locule. It dehisces from two indehiscent, single-seeded mericarps
attached with carpophore, e.g., Umbelliferae.
11. Cypsela A dry, indehiscent achene type of fruit derived from unilocular, inferior ovary, e.g.
Helianthus.
12. Drupe A fleshy fruit with a stony endocarp and edible mesocarp, e.g. mango, Prunus.
13. Druplet A small drupe, e.g. Rubus.
14. Etaerio of Achenes Also called achenecetum, it is an aggregation of achenes, e.g.
Ranunculus.
15. Etaerio of Berries Also called baccacetum, it is an aggregation of berries, e.g.
Artabotrys.
16. Etaerio of Drupes Also called drupecetum, it is an aggregation of drupes, e.g. Rubus.
17. Etaerio of Follicles Also called follicetum, it is an aggregation of follicles, e.g.
Delphinium.
18. Etaerio of Samaras Also called samaracetum, it is an aggregation of samaras, e.g.
Liriodendron.
19. Follicle A dry, dehiscent fruit derived from a monocarpellary, superior ovary, and dehisces
only along one suture, e.g. Delphinium.
20. Hesperidium A fleshy, thick-skinned berry with hard and leathery pericarp. It is derived
from a polycarpellary, multilocular, superior ovary. Bulk of this fruit is derived from glandular
hairs, e.g. Citrus.
1
550 Terms of Plant Description
Achene
Berry
(Mirabilis) Cacervulus Capsule
(Tomato)
(Althea) (Papaver)
Epicarp
Scar of Style
Mesocarp
Endocarp
Seed
Caryopsis
(Zea mays) Cremocarp
Cypsela
(Coriandrum)
(Helianthus) Drupe
(Mango)
Fig. 20.17 Types of fruit (arranged alphabetically).

207
208
Persistent
style
Etaerio of
Etaerio of Etaerio of
Achenes Etaerio of
Berries Drupes
(Naravelia) Follicles
(Artabotrys) (Rubus) (Michelia)
Follicle Hesperidium Legume Lomentum

(Delphinium) (Orange) (Pisum) (Acacia)

Plant Taxonomy
Pome Regma
Nut (Apple) (Ricinus)
(Anacardium)
Samara Siliqua
Silicula (Elm) (Brassica)
(Iberis)
Sorosls
(Morus)
Syconus Utricle
Sorosis (Ficus) (Chenopodium)
(Pineapple)
21. Legume A dry, dehiscent fruit derived from monocarpellary, superior ovary with marginal
placentation. It dehisces along two sutures, e.g. Pisum.
22. Lomentum A dry, dehiscent legume that separates transversely between seed sections, e.g.
Acacia.
23. Nut A dry, indehiscent, one-seeded fruit with a hard pericarp. It is derived from unilocular
ovary, e.g. Anacardium.
24. Nutlet A small nut.
25. Pepo A berry with a leathery nonseptate rind, developing from tricarpellary, syncarpous,
inferior ovary with parietal placentation, e.g. Cucurbitaceae.
26. Pome A fleshy fruit surrounded by the fleshy thalamus and developing from a two- or
more-celled, syncarpous, inferior ovary, e.g. apple.
210 Plant Taxonomy
27. Pyrene Fleshy fruit with each seed covered by a bony endocarp, e.g. Ilex.
28. Regma Dry, schizocarpic fruit derived from tricarpellary, syncarpous, superior, trilocular
ovary, and bears many spinous tubercles, e.g. Ricinus.
29. Samara Dry, indehiscent, simple fruit derived from bicarpellary, syncarpous ovary. Its
pericarp is winged, e.g. Elm.
30. Silicula Dry, dehiscent fruit derived from two or more carpels. It dehisces along two sutures
and leaves a persistent partition after dehiscence. It is as broad as, or even broader, than long,
e.g. Iberis.
31. Siliqua Dry, dehiscent fruit derived from bicarpellary, syncarpous ovary with parietal pla-
centation and a false septum. It is longer than broad, e.g. Brassica.
32. Sorosis A multiple fruit developing from the spadix or spike in which the flowers usually
fuse by their succulent sepals, and axis bearing them becomes woody or fleshy forming a
compact mass, e.g. pineapple.
33. Syconus A multiple fruit derived from hypanthodium type of inflorescence. Here the
achenes develop on the inside of a hollowed-out fleshy receptacle, e.g. Ficus.
34. Utricle Dry, indehiscent, small, bladder-like, one-seeded fruit, e.g. Chenopodium.

1. Differentiate between the following:
(a) Shrub and tree (b) Adventitious root and tap root
(c) Fusiform root and napiform root (d) Cladode and corm
(e) Phyllode and phylloclade (f) Tendril and thorn
(g) Bract and bracteole (h) Calyx and epicalyx
2. Make diagrams of the following leaf apices and explain each of them only in one
sentence:
(a) Acute (b) Acuminate
(c) Mucronate (d) Obtuse
3. Explain the following types of leaf margins using only one sentence for each of them:
(a) Dentate (b) Entire
(c) Revolute (d) Serrate.
4. How will you differentiate between alternate, opposite and whorled type of phyllotaxies of
leaf?
5. With the help of diagrams, differentiate between parallel and reticulate venations. What is
the difference between unicostate and multicostate venations?
6. Explain briefly following types of inflorescence:
(a) Corymb (b) Capitulum
(c) Spadix (d) Verticillaster.
7. What do you mean by aestivation? Explain various types of aestivations with the help of
suitable diagrams.
8. With reference to androecium, explain the following terms:
(i) Basifixed (ii) Didynamous
(iii) Monadelphous (iv) Obdiplostemonous
(v) Syngenesious (vi) Staminode
9. Explain major types of placentations using suitable diagrams.
10. Give one example each of the plants bearing the following types of fruits:
(a) Berry (b) Caryopsis
(c) Hesperidium (d) Sorosis
Suggested Reading
Jones, S.B. Jr. and A.E. Luchsinger, 1987, Plant Systematics (2nd ed.), McGraw-Hill, New York.
Lawrence, G.H.M., 1951, Taxonomy of Vascular Plants, Macmillan Company, New York.
Sharma, O.P., 1993, A Manual of Practical Botany Vol. II. (5th ed.), Pragati Prakashan, Meerut.
Sugden, A., 1984, Longman Illustrated Dictionary of Botany, Longman, U.K.
C
H
A
P
T
E
R
SELECTED
FAMILIES OF
DICOTYLEDONS 21
Various workers have divided Dicotyledons into varying number of families. Bentham and Hooker
(1862–1883) discussed 163 families under Dicotyledons in their Genera Plantarum while Armen
Takhtajan (1964) divided Magnoliatae (or Dicotyledones) into 361 families in his book entitled
Flowering Plants: Origin and Dispersal. Arthur Cronquist (1981) in his treatment An Integrated
System of Classification of Flowering Plants, included 318 families under class Magnoliopsida
(=Dicots) whereas Robert F. Thorne (1983) treated Dicotyledoneae (=Annonidae) as a subclass of
class Angiospermae (=Annonopsida) and discussed 297 families under this subclass. Thorne’s system
of classification is published in Nordic Journal of Botany (1983).
The present chapter deals with the study of the taxonomic characters of 73 selected families of
Dicotyledons. Those that have been selected were chosen because of their common availability,
unusual characters, great economic importance, large number of recorded species, and/or showing
typical representation of any particular order or group.
Compilation of characters of each family has been done from the available literature, including
Lawrence (1951), Hutchinson (1973), Willis (1973), Heywood (1978), Radford (1986), and Jones and
Luchsinger (1987). The arrangement of families in this chapter follows that of Bentham and Hooker
(1862–1883) because this old but natural system of classification still suits best to the students as
well as teachers, specially in the laboratory. However, the position of each family in some widely
accepted systems of classification (Engler and Prantl, 1887–1915; Bessey, 1915; Takhtajan, 1969;
Hutchinson, 1973; Cronquist, 1981; and Thorne, 1983) is also discussed separately under systematics
and phylogeny. Economic importance of each family is also discussed in some detail.
21.1 DICOTYLEDONS
The dicotyledons include all those angiosperms in which the embryo possesses two cotyledons,
leaves with reticulate venation and vascular bundles are open and arranged in one or more rings.
These plants have secondary thickenings in the stems. Due to the presence of cambium, these plants
may be either woody or herbaceous. Dicotyledons usually have pentamerous flowers. They possess
a persistent primary root that develops into a tap root.
Selected Families of Dicotyledons 213
Bentham and Hooker (1862–1883) divided the class Dicotyledons into three subclasses, viz.
Polypetalae, Gamopetalae and Monochlamydeae.
21.2 POLYPETALAE
The members of subclass Polypetalae contain flowers with free petals, and their perianth is usually
in two whorls i.e. calyx and corolla. Polypetalae has been divided into 3 series, viz. Thalamiflorae,
Disciflorae and Calyciflorae.
Series Thalamiflorae is characterised by (i) usually distinct sepals free from ovary, (ii) pres-
ence of many stamens, (iii) hypogynous flowers, (iv) superior ovary, and (v) absence of disc.
Thalamiflorae includes 6 cohorts (=orders) and 34 orders (=families). Six cohorts are Ranales,
Parietales, Polygalineae, Caryophyllineae, Guttiferales and Malvales.
Series Disciflorae is characterised by (i) distinct or united sepals, free or adnate to ovary, (ii) pres-
ence of disc, (iii) stamens hypogynous, usually definite, (iv) superior ovary. Disciflorae includes 4
cohorts (=orders) and 23 orders (=families). The cohorts are Geraniales, Olacales, Celastrales and
Sapindales.
Series Calyciflorae is characterised by (i) usually inferior ovary, (ii) united or rarely free sepals,
and (iii) flowers perigynous or epigynous. It includes 5 cohorts (=orders) and 27 orders (=families).
The cohorts are Rosales, Myrtales, Passiflorales, Ficoidales and Umbellales.
Few general characters of different orders alongwith some detailed description of some representa-
tive families are discussed in the forthcoming text.
21.3 RANALES
21.3.1 General Characteristics
1. Flowers usually with numerous and indefinite number of stamens, and only rarely the stamens
number is definite.
2. The gynoecium apocarpous and multicarpellate, i.e. composed of many unilocular unicarpel-
late pistils.
3. Embryo minute and seeds endospermic.
4. Floral parts typically spirally arranged, numerous and distinct.
5. Perianth often not clearly differentiated into calyx and corolla.
Bentham and Hooker included 8 orders (=families) under Ranales. These are Ranunculaceae,
Mangnoliaceae, Annonaceae, Berberidaceae, Nymphaeaceae, Menispermaceae, Dilleniaceae and
Calycanthaceae.
Engler and Diels, however, divided the order Ranales into 4 suborders containing 19 families, as
under:
Suborder Nymphaeineae: Nymphaeaceae, Ceratophyllaceae
Suborder Trochodendrineae: Trochodendraceae, Cercidiphyllaceae
Suborder Ranunculineae: Ranunculaceae, Lardizabalaceae, Berberidaceae,
Menispermaceae.
214 Plant Taxonomy
Suborder Magnoliineae: Magnoliaceae, Himantandraceae, Calycanthaceae, Lactoridaceae,

Annonaceae, Eupomatiaceae, Myristicaceae, Gomortegaceae,
Monimiaceae, Lauraceae, Hernandiaceae.
Only Magnoliaceae, Annonaceae, Ranunculaceae and Nymphaeaceae are discussed in this text.
21.4 MAGNOLIACEAE MAGNOLIA FAMILY

21.4.1 Systematic Position
Polypetalae, Thalamiflorae, Ranales.
21.4.2 Field Recognition

Trees or shrubs; leaves simple, alternate; stipules leave a circular stipular scar; flowers large, acti-
nomorphic; perianth parts 6–18; stamens many; apocarpous.
21.4.3 Selected Indian Genera

Magnolia, Michelia, Liriodendron, Aclimandra, Talauma, Manglietia and Schizandra.
21.4.4 Size, Distribution, and General Information

Named after a French botanist Pierre Magnol, Magnoliaceae is a family of about 12 genera and 230
species, mainly distributed in the warm tamperate regions of the Northern Hemisphere. The species
are common in eastern Asia and eastern North America, and found also in Brazil, West Indies, and
Malaysia. 36 species of this family have so far been reported from different parts of India.
21.4.5 Description of the Family

General Habit Deciduous or evergreen, often aromatic shrubs or trees; Liriodendron tulipifera
reaches 20 to 65 m in height; Michelia and Magnolia are medium-sized trees or shrubs; Schizandra
is a climber.
Root Tap root, branched.
Stem Aerial, woody, erect or climbing, branched; parenchyma contains oil glands.
Leaves Alternate, large, simple, entire, petiolate, pinnately-veined; stipulate, stipules large, protect
young buds and leave a circular scar on falling; stipules absent in Wintereae; oil glands present.
Inflorescence Flowers usually solitary, either axillary (Michelia), or terminal (Magnolia); flowers
crowded near the tips of branches in Illicium.
Flowers Large, showy, bracteate (Magnolia, Michelia) or ebracteate, pedicellate, bisexual but rarely
unisexual in Kmeria; actinomorphic; hypogynous; floral parts arranged spirally on the elongated
floral axis; some Drimys species have unisexual flowers while others have bisexual flowers.
Perianth Except in Illicium and some species of Magnolia, Michelia and Drimys, the perianth
is undifferentiated into calyx and corolla. Perianth spiral to cyclic, in 3 or more series of 6 to 18
parts; polyphyllous. In Drimys sepals fused while petals

free (Fig. 21.1). Perianth lobes are arranged in whorls of
threes, valvate or imbricate in each whorl; 9–12 perianth
lobes in Magnolia while 6–15 in Michelia. In Euptelea,
perianth is absent.
Androecium Stamens numerous, free, spirally arranged
on the basal part of floral axis; anthers long, dithecous,
basifixed or adnate; introrse but extrorse in Liriodendron;
dehiscence longitudinal; connective prolonged.
Gynoecium Carpels numerous, apocarpous (free), arranged
spirally on the floral axis above the stamens; superior, uni-
locular, one or more ovules on a ventral suture; placenta- Fig. 21.1 Floral diagram of Drimys.
tion parietal; style short, one; stigma one. In Zygogynum
and Pachylarnax, the carpels are rarely united.
Fruits and Seeds Fruit of aggregate type. It may be a follicle (Magnolia), samara (Liriodendron), or
rarely berry (Schizandra). Seed often suspended within the follicle by a thread-like funiculus, has a
minute embryo in a large endosperm.
Pollination and Dispersal Pollination is entomophilous; dispersal takes place by those animals (e.g.
birds, monkeys, etc.) which carry off the fruits.
General Floral Formula ≈, (rarely unisexual), P9 – μ, A μ, G μ.
21.4.6 Economic Importance

• The Magnoliaceae are important as common ornamentals and as a source of wood.
• Magnolia species are commonly cultivated in gardens. M. grandiflora (Dulee Champa,
or ‘Bari Champa’), M. fuscata (Chidi Champa), M. campbelli (‘Lal Champa’), M. pumila
(‘Jauhre Champa’), M. pterocarpa and M. stellata are known for their sweet, scented, beau-
tiful flowers. Wood of Magnolia acuminata is used in making toys, tea boxes and quality
furniture.
• Michelia champaca (‘Champa’) is a common garden ornamental plant, and is also the source
of a volatile oil which is widely used in perfumery and cosmetics. Its flowers and fruits are
used to cure kidney troubles and gonorrhoea. M. fuscata flowers yield an essential oil which
is used as hair oil. M. alba, M. excelsa and M. nilogrica are other ornamental species with
sweet-scented flowers.
• Liriodendron tulipifera provides a tall graceful posture and bears beautiful scented flowers.
It yields good timber for furniture.
• Talauma phellocarpa trees yield good quality timber used in making tea boxes and
furniture.
• Manglietia hookeri provides good furniture wood.
216 Plant Taxonomy
• Wood of Pachylarnax and Aclimandra cathcartii is also useful in making indoor

furniture.
• Illicium verum yields an oil useful in colic disorders. I. griffithii fruits are used as condi-
ment for flavoring curries.
• Drimys winteri bark is used as an astringent and stimulant.
• Schizandra grandiflora fruits are edible.
21.4.7 Systematics and Phylogeny (Affinities)

Family Magnoliaceae is divided into three tribes (Wintereae, Magnolieae and Schizandreae) by
Bentham and Hooker (1867), three subfamilies (Magnolioideae, Illicioideae and Schizandroideae) by
Rendel (1938), and only into two tribes (Magnolieae and Liriodendreae) by Jones and Luchsinger
(1987). Several taxonomists treated families, such as Eupteleaceae, Illiciaceae, Schizandraceae,
Trochodendraceae and Winteraceae as belonging to family Magnoliaceae. But Hutchinson (1959)
has treated them as independent families, except Eupteleaceae, which was merged by him under
Schizandraceae. Takhtajan (1969) also treated them as independent families.
Takhtajan (1969), Hutchinson (1973), Cronquist (1981) and several other taxonomists opine that
Magnoliaceae is the most primitive family of angiosperms.
21.4.8 Description of Some Important Plants

1. Michelia champaca L. (Vern. “Champa”; Fig. 21.2)
Habit: A small evergreen tree. Root: Branched, tap root. Stem: Erect, aerial, branched, woody. Leaf:
Simple, alternate; stipulate, stipule convolute; petiolate, ovate, entire, acute, unicostate reticulate.
Inflorescence: Solitary axillary. Flower: Bracteate, pedicellate, complete, hermaphrodite, actinomor-
phic, hypogynous, fragrant, pale yellow. Perianth: Usually 9 perianth leaves arranged in three whorls
of three each; either outer 3 leaves are sepalloid and inner 6 lobes are petalloid, or all the 9 perianth
lobes are petalloid. Androecium: Stamens numerous, free, spirally arranged on a conically elongate
receptacle, filaments short, basifixed, dithecous, extrorse. Gynoecium: Multicarpellary, apocarpous,
carpels arranged spirally over the conical receptacle, superior, unilocular, many ovules, marginal
placentation; style curved, stigma beaked. Fruit: Etaerio of follicles.
Floral Formulae: Br, ≈, , K3, C3+3, A μ, G μ; or Br, ≈, , P3+3+3, A μ, G μ.
2. Magnolia grandiflora L. (Vern. “Dulee Champa”)
Habit: A medium-sized, evergreen tree. Root: Branched, tap root. Stem: Aerial, erect, branched, solid,
woody. Leaf: Alternate, simple; stipulate, stipules large and cover the bud; lanceolate, entire, acute,
unicostate reticulate. Inflorescence: Solitary terminal. Flower: Ebracteate, complete, hermaphrodite,
actinomorphic, hypogynous, fragrant, white. Perianth: 9 lobes, in three whorls of three each, valvate
in each whorl, white. Androecium: Numerous stamens, polyandrous, arranged spirally on receptacle,
dithecous, filament small, anther lobes cone-shaped, connective indistinct; introrse, dehiscence lon-
gitudinal. Gynoecium: Same as in Michelia. Fruit: Same as in Michelia.
Floral Formula: Ebr, ≈, , P3+3+3, A μ, G μ.
style perianth
leaves
stigma
ovary
carpel
stamen
pedicel
stigma
A Carpel
L.S. Flower
style
ovule
ovary
Carpel
flower
leaf
stem
Floral Diagram Flowering Branch
Fig. 21.2 Michelia champaca L.
21.5 ANNONACEAE CUSTARD APPLE FAMILY


Woody shrubs, trees or climbers; wood aromatic; leaves two-ranked, without stipules; perianth
3 + 3 + 3; stamens numerous, rarely few; gynoecium polycarpellary, apocarpous, superior ovary.
218 Plant Taxonomy

Annona squamosa (Vern. “Sharifa”), Cananga odorata (Ylang-ylang), Artabotrys odoratissimus.

This largest family of order Magnoliales (130 genera and 2300 species, Jones and Luchsinger, 1987)
is widely distributed in the Old World Tropics. Over 25 genera and 200 species have so far been
reported from India, mostly from its peninsular region. Annona is represented by over 120 species,
and Xylopia by about 150 species.

General Habit Usually aromatic trees or shrubs, rarely liana-like, or woody climbers, e.g. species of
Oxymitra. Artabotrys is a hook climber. Bark, leaves and floral parts contain oil ducts.
Root Extensively branched tap roots.
Stem Woody, branched, erect, rarely climbing, aerial, hard.
Leaves Simple, alternate, two-ranked, exstipulate, petiolate; apex acute, obtuse or even mucronate
(Annona mucronata); ovate, obovate or lanceolate; coriaceous, glabrous; unicostate reticulate; gland-
dotted, and so aromatic.
Inflorescence Solitary axillary or terminal, or leaf opposed; sometimes cauliflorous, i.e. flowers
occur on stem (e.g. Polyalthia fragrans).
Flower Bracteate or ebracteate, ebracteolate, pedicellate, complete, hermaphrodite, rarely unisexual
(Stelechocarpus); actinomorphic, rarely zygomorphic (Monodora); hypogynous.
Calyx Sepals 3, or sometimes 6 arranged in two whorls of three each; polysepalous, or rarely basally
connate; valvate, rarely imbricate.
Corolla Generally 6 petals, arranged in two whorls of three each, sometimes only 3 petals; poly-
petalous; valvate or slightly imbricate.
Sometimes, sepals and petals are alike, and thus represent the perianth.
Androecium Stamens numerous, polyandrous; spirally arranged on large convex receptacle; fila-
ments short and thick; connective continues beyond the anther and produces a dilated head; adnate;
extrorse.
Gynoecium Carpels numerous, arranged spirally on the raised receptacle (Polyalthia, Artabotrys);
only 1–3 carpels in Cyathocalyx; apocarpous; superior, unilocular, one to many anatropous ovules
in the locule; parietal or sometimes basal or marginal placentation; style short or absent; stigma
sessile, simple or trilobed (Tetrameranthus).
Fruits and Seeds Fruit an etaerio of berries; in Annona, the berries coalesce with fleshy receptacle;
seeds large, endospermous.
Pollination and Dispersal Pollination generally by flies; dispersal by birds, bats or mammals.
Floral Formulae Br or Ebr, ≈, , K3 or 3+3, C3+3 or 3, A μ, G μ
Or, Br or Ebr, ≈, , P3+3+3, A μ, G μ.

• Annona squamosa (Vern. “Sharifa” or “Sitaphal”) is cultivated in Assam, West Bengal,
Maharashtra, etc. for its edible fruits. Its leaves and seeds have insecticidal properties.
A. reticulata (Vern. “Ramphal”), A. muricata (Vern “Maniphal”), A. cherimola (Vern.
Hanumanphal), A. diversifolia are all grown for their edible fruits.
• Artabotrys odoratissimus (“climbing ylang-ylang”) yields an ethereal oil and also cultivated
for its scented flowers.
• Polyalthia longifolia and P. pendula (Vern. Ashok) are grown for their beautiful green
drooping branches on road sides and in beautiful residential buildings. P. longifolia wood
is used for making pencils, boxes, match box industry, etc.
• Asimina triloba is cultivated for its edible fruits in America.
• Xylopia aethiopica is the source of Guinea pepper. X. parvifolia wood is used in making
plywood.
• Cananga odorata is cultivated as an ornamental tree because of its sweet-smelling flowers.
An oil, obtained from its flowers, is used in perfume industry.
• Unona pannosa yields a strong fibre used for paper making and cordage.
• Monodora myristica is cultivated for its beautiful flowers. Its seeds contain oil used as a
nutmeg substitute.
• Uraria narum roots yield an oil used as a medicine.
• Bark of Goniothalamus yields strong fibre.

The resemblances and differences between Magnoliaceae and Annonaceae indicate a close relation-
ship between the two families. The two families resemble with each other in possessing (i) large
solitary flowers having perianth leaves in more than one whorl, (ii) spirally arranged stamens,
(iii) polycarpellary, apocarpous condition, (iv) large aggregate fruits, (v) seeds with large endosperm
and small embryo, (vi) anatropous ovules, (vii) oil-containing cells; (viii) woody stems. etc. On the
other hand, Annonaceae differs from Magnoliaceae in possessing (i) exstipulate leaves, (ii) extrorse
stamens, (iii) ruminant endosperm, and (iv) sarcotesta in the seeds.
The above-mentioned resemblances and differences between Annonaceae and Magnoliaceae
indicate that both of them are closely related, and have perhaps been derived from the same ances-
tral stock. Therefore, Hutchinson’s (1973) placing them in two separate orders (Magnoliaceae in
Magnoliales and Annonaceae in Annonales) is not advisable. Cronquist (1981) placed both of them
under Magnoliales, and Thorne (1983) under suborder Annonineae of order Annonales, and this
appears justified.
Because of some characters, Annonaceae are more advanced than Magnoliaceae.
Engler and Prantl (1887–1899) divided family Annonaceae into five tribes, viz. Uvarieae, Miluseae,
Hexalobieae, Xylopeae, and Monodoreae.
220 Plant Taxonomy
21.5.8 Description of Some Important Plants

1. Artabotrys odoratissimus syn. A. uncinatus (Vern. Climbing Ylang-Ylang or Kantali Champa; Fig. 21.3)
Habit: Woody climber, climbing with the help of hooks developed on flower pedicels. Root: Branched,
tap roots. Stem: Aerial, woody, branched, climbing, aromatic. Leaf: Simple, alternate, exstipulate,
petiolate, elliptical, entire, acute apex, unicostate reticulate; oil ducts are present. Inflorescence:
Solitary axillary, or rarely in groups. Flower: Bracteate, pedicellate, pedicel bears a hook; com-
plete, hermaphrodite, actinomorphic, trimerous, hypogynous, spirocyclic. Perianth: Perianth lobes 9,
arranged in 3 whorls of 3 each; sometimes, outer whorl of 3 lobes is green sepalloid, free, valvate,
and inner 6 lobes of two whorls are petalloid, valvate, or slightly imbricate; perianth lobes are
sweet-scented and gland-dotted. Androecium: Stamens numerous, polyandrous, spirally arranged
on the receptacle; filaments short, connective prolonged beyond the anther; extrorse. Gynoecium:
Multicarpellary, apocarpous, carpels spirally arranged on receptacle; ovary superior, unilocular, one
to many anatropous ovules in each locule; marginal placentation. Fruit: Etaerio of berries. Seed:
Large, endospermic.
Floral Formulae: Br, ≈, , P3+3+3, A μ, G μ
Or, Br, ≈, , K3, C3+3, A μ, G μ.
prolonged
carpels connective
anther
stamen
inner perianth lobe filament
An Enlarged Stamen
outer perianth
lobe
A Dissected Flower
A Flower
flower
stem
Fruits
Flowering Branch
Floral Diagram
Fig. 21.3 Artabotrys odoratissimus R. Br.
2. Annona squamosa (Vern. “Sharifa”)

Habit: A tall shrub or small tree. Stem: Erect, woody, branched. Leaf: Simple, alternate, exstipulate,
oblanceolate, entire, obtuse. Inflorescence: Axillary, 2 or more flowers developing in the axis of
leaf. Flower: Bracteate, pedicellate, hermaphrodite, actinomorphic, hypogynous, spirocyclic. Calyx:

Sepals 3, poly- or gamosepalous, valvate. Corolla: Petals 3, or 6, arranged in two whorls of 3 each;
polypetalous, valvate. Androecium: Anthers appendaged; other details similar to Artabotrys odorat-
issimus. Gynoecium: Multicarpellary, apocarpous; carpels arranged spirally on receptacle; superior,
unilocular, only one ovule in the locule, basal placentation. Fruit: Etaerio of berries, or of achenes.
Seed: Endospermic.
Floral Formula: Br, ≈, , K3 or (3), C3 or 3 + 3, A μ, G μ.
21.6 RANUNCULACEAE BUTTERCUP FAMILY


Herbs; leaves often divided or palmately compound with sheathing leaf bases; stamens numerous
and spirally arranged; carpels numerous, distinct, spirally arranged; unilocular.

Ranunculus, Delphinium, Aconitum, Actaea, Aquilegia, Anemone, Caltha, Clematis, Helleborus,
Nigella, Paeonia, Thalictrum.

This family is represented by about 50 genera and 1900 species, chiefly distributed in the temperate
regions of the Northern Hemisphere. Eastern Asia and eastern North America are the regions of
their most common occurrence. The common genera, of which over 150 or more species have so
far been reported, include Ranunculus (400), Aconitum (300), Delphinium (250), Clematis (250),
Anemone (150) and Thalictrum (150). Some species occur in Arctic and Alpine zones. However,
they are rare in tropics. 20 genera and over 165 species of this family have so far been reported
from India. Plants of this family flower mainly during winters in our country.

General Habit Generally annual or perennial herbs, rarely shrubs (Xanthorhiza) or vines (Clematis,
Naravelia); perennials perennate by means of rhizome (Cimicifuga, Helleborus) or tuberous roots
(Aconitum); some species are aquatic herbs (Ranunculus aquatilis).
Root Tap root as well as adventitious roots; tuberous roots of Aconitum, Paeonia and Ranunculus
store food, swell and form tuber-like structures.
Stem Aerial as well as underground; herbaceous (Ranunculus); woody (Paeonia); climbing
(Clematis); branched.
Leaves Usually basal and cauline; often palmately divided with sheathing leaf bases; usually
exstipulate, but stipulate in Trollius, Caltha, Thalictrum and Ranunculus; mostly alternate, rarely
222 Plant Taxonomy
opposite (Clematis); simple (Ranunculus), pinnately compound (Clematis), decompound (Thalictrum),

or finely dissected into fine segments (Delphinium ajacis); aquatic species of Ranunculus show
heterophylly.
Inflorescence Variously formed, from solitary (Nigella) to paniculate (Thalictrum), or in racemes
(Delphinium) or cymes; a single terminal flower is produced in Anemone and Eranthis, but more
often a cymose branching is seen.
Flower Bracteate (Clematis) or ebracteate (Anemone), bracteolate (Delphinium) or ebracteolate
(Ranunculus), usually bisexual but unisexual in some species of Thalictrum; usually actinomor-
phic but zygomorphic in Delphinium and Aconitum; hypogynous; spurred in Delphinium and
Aquilegia.
Calyx 3 to many sepals, usually 5 (Ranunculus), polysepalous, valvate or imbricate; sepals are
petaloid in Aconitum and Delphinium and their posterior sepal is spurred. In Nigella and Anemone
there is sometimes an involucre of green leaves below the flower, usually alternate with the calyx.
Corolla Usually 5 petals, polypetalous; nectariferous glands often present; sometimes petals are
many or absent (Caltha); posterior petals are spurred.
In genera such as Paeonia (Fig. 21.4), Clematis (Fig. 21.5), Actaea and Helleborus, perianth is
present i.e. there is no distinction between sepals and petals; innermost perianth leaves often bear
nectaries (honey-leaves) or they are reduced to small nectariferous scales.
Androecium Stamens 5 to many, free, spirally arranged; anthers 2-celled, adnate, extrorse and
longitudinally dehiscent; sometimes staminodes are present (Aquilegia); some outer stamens may
be converted into honey-leaves (Helleborus); from 80 to 150 stamens in Caltha; stamens bright
coloured in Thalictrum.
Gynoecium Usually numerous, free carpels with either one basal ovule or several anatropous ovules
showing marginal placentation; ovary superior, unilocular; style 1; stigma 1; in Actaea there is only
1 carpel; in Nigella carpels are united, ovary pentalocular and placentation axile; in Helleborus
carpels are only basally connate.
Fruits and Seeds Fruit may be an etaerio of follicles (Aconitum), etaerio of achenes (Ranunculus),
a berry (Actaea), or capsule (Nigella); seeds with minute embryo and oily endosperm; elaisome (an
appendage which contains oily substance attractive to ants) is present on the seeds of some species
of Helleborus.
On the basis of the type of fruit and number of carpels, family Ranunculaceae is divided into
two subfamilies:
1. Helleboroideae: Carpels with more than 1 ovule; fruit a berry or follicle.
2. Ranunculoideae: Carpels with 1 ovule; fruit etaerio of achenes, rarely a berry.
Pollination and Dispersal The flowers are usually protandrous in Ranunculaceae and therefore well-
suited for insect pollination. Insects are also attracted because of the presence of nectar, petaloid
calyx, conspicuous corolla and stamens. However, some species of Thalictrum are wind-pollinated,
e.g. T. minus. Dispersal takes place by animals or birds (Ranunculus) or by wind (Anemone,
Clematis).
General Floral Formula Br or Ebr, ≈ or , , K5 or K3 – μ, C5 or P4 – 6, A5–μ, Gμ or 1 – μ.
Fig. 21.4 Floral diagram of Paeonia. Fig. 21.5 Floral diagram of Clematis.

Ranunculaceae are important as ornamental, and as drug plants. They are mostly poisonous owing
to the presence of alkaloids.
(a) Ornamentals Many Ranunculaceae have showy flowers and are grown as ornamentals
in our gardens. Some of such ornamentals along with their common name in parenthesis
are Aconitum napellus (Aconite), Anemone parviflora (Wind flower), Aquilegia coerulea
(Columbine), Caltha palustris (Marsh marigold), Clematis virginiana (Traveller’s joy),
Clematis vitalba (Virgins-bower), Delphinium ajacis (Larkspur), Helleborus (Christmas
rose), Nigella damascena (Love-in-a-mist), Paeonia emodi (Peony), Ranunculus (Butter-
cup, e.g. R. asiaticus, R. muricatus, R. repens etc.), and Thalictrum alpinum (Meadow-rue).
Species of Anemone, Clematis, Delphinium, Ranunculus and Thalictrum are available in a
wide range of colours and forms.
(b) Medicinal Uses Large number of plants of this family are of medicinal value, of which
some are undermentioned:
1. Aconitum napellus roots are the source of aconite, used as a nerve sedative, and for
rheumatism. Its dried roots are used to cure mild heart palpitation and gastritis. Deadly
poisonous alkaloid, obtained from A. ferox, may cause death but is useful in snake
bites.
2. Actaea spicata roots are used as a nerve sedative.
3. Adonis aestivatis and A. scorbiculata roots are used as heart stimulant.
4. Anemone pulsatilla is the source of the homoeopathic drug ‘pulsatilla’ used to cure
nervous disorders in women during menstrual periods.
5. Clematis triloba leaves are used to cure snake bites.
6. Coptis teeta roots are used in brain and liver disorders as well as in toothache and sore
eyes.
224 Plant Taxonomy
7. Delphinium ajacis and D. elatum seeds are insecticidal while roots of D. denudatum
are effective in toothache. Seeds, leaves and roots of D. zatil are used to cure jaundice
and spleen. D. vestitum seeds are useful in cardiac disorders.
8. Helleborus niger flowers are given in the form of syrup to mad persons. This species
is also useful in chronic skin infections.
9. Nigella sativa (vern. Kalonji or Kala Zeera) seeds are used as common condiment as
well as in the treatment of cough, asthma and fever.
10. Paeonia officinalis tubers are used to cure colic and uterus disorders.
11. Ranunculus muricatus and R. arvensis juice is used in irregular fever. R. aquatilis and
R. arvensis leaves and flowers are used to cure asthma and rheumatism. R. ficaria is
used to cure piles while R. falcatus is poisonous and causes skin blisters.
12. Thalictrum foliolosum roots are used as diuretic, purgative and tonic.

Majority of the phylogenetists believe that Ranunculaceae is one of the most primitive families of
dicotyledons because of the primitive features, such as (i) presence of scattered vascular bundles in
the stem in some of its taxa, such as Actaea and Thalictrum, (ii) spiral arrangement of foliage and
floral leaves, (iii) bisexual, hypogynous, actinomorphic flowers with apocarpous gynoecium, (iv)
seeds with small embryo and oily endosperm, and (v) anatropous ovules. On the other hand, this
family also shows several advanced characters such as (i) presence of unisexual and zygomorphic
flowers in some genera, (ii) fusion of carpels (Nigella), and (iii) achene type of fruits (Ranunculus).
Several phylogenetists believe that monocots have originated from dicots, and the point of their origin
being Ranunculaceae or some allied groups of Ranales.
Ranunculaceae is divided into five tribes (Anemoneae, Clematideae, Helleboreae, Paeonieae and
Ranunculeae) by Bentham and Hooker (1862–1883). However, Engler and Prantl (1887–1899) recogn-
ised only three tribes, viz. Anemoneae, Helleboreae and Paeonieae. Hutchinson (1973) separated tribe
Helleboreae from Ranunculaceae and raised it to the rank of an independent family Helleboraceae.
Hutchinson (1973), Takhtajan (1980) and Cronquist (1981) have also raised tribe Paeonieae to the
rank of a family Paeoniaceae with the genus Paeonia as its sole representative.
Jones and Luchsinger (1987) have stated that in Ranunculaceae, the perianth varies greatly and
is of importance in delimiting genera in this family.
Hickey and King (1988) divided family Ranunculaceae into 2 sub-families, i.e. Helleboroideae
and Ranunculoideae. Helleboroideae is again divided into 2 tribes, i.e. Helleboreae and Delphinieae
while Ranunculoideae into 3 tribes, i.e. Ranunculeae, Anemoneae and Clematideae.
21.6.8 Description of Some Common Plants

1. Ranunculus sceleratus L. (Vern. Celery-leaved Crow-foot; Fig. 21.6)
Habit: Annual herb. Stem: Branched, erect, glabrous and solid. Leaf: Simple, alternate, petiolate, exs-
tipulate, leaf base sheathing, divided further into obovate or cuneate segments, multicostate reticulate.
petal stamen
sepal carpels flowers
anther lobe
receptacle pedicel connective

L.S. Flower
filament fruit
stem
A stamen
leaf
roots
Floral Diagram
Flowering Plant
Fig. 21.6 Ranunculus sceleratus L.
Inflorescence: Cymose (dichasial cyme). Flower: Bracteate, bracteolate, pedicellate, complete, her-
maphrodite, actinomorphic, pentamerous, hypogynous, spirocyclic. Calyx: 5 sepals, polysepalous,
imbricate or quincuncial, petaloid, reflexed from the base. Corolla: 5 petals, polypetalous, imbri-
cate, bright yellow, a nectary is present at the base of each petal. Androecium: Numerous stamens,
polyandrous, spirally arranged; anthers dithecous, basifixed, extrorse. Gynoecium: Multicarpellary,
apocarpous; each carpel superior, unilocular; one ovule in each locule; basal placentation; carpels
arranged spirally over an oblong receptacle; style reduced; stigma fimbriate. Fruit: Etaerio of achenes.
Seed: Endospermic.
Floral Formula: Br, Brl, ≈, , K5, C5, A μ, G μ.
226 Plant Taxonomy
2. Delphinium ajacis Auct. (D. ambiguum L., Consolida ambigua (L.) Ball. and Heyw; Vern. Larkspur;
Fig. 21.7)
Habit: Cultivated, annual herb. Stem: Erect, branched, solid, green, hairy. Leaf: Simple or decom-
pound, alternate, exstipulate, deeply dissected; each lobe is linear to elliptical, smooth, with acute or
postero-lateral petal
lateral sepal
posterior sepal
antero-lateral
petal
anterior-sepal
seed
spur
lateral sepal
A Flower (front view)
locule
ovule
stamen Fruit
ovary
wall
T.S. ovary ovary stigma

sepal
spur
petal
pedicel
bracteole
L.S. Flower
stem
flower
anther lobe
connective
filament
Floral Diagram A Stamen Flowering Branch
Fig. 21.7 Delphinium ajacis L.

acuminate apex. Inflorescence: Axillary raceme of 4 to 16 flowers. Flower: Bracteate, bracteolate,

pedicellate, complete, hermaphrodite, zygomorphic, spurred posteriorly, pentamerous, hypogynous;
bright blue, pink or white. Calyx: 5 sepals, polysepalous, petaloid; posterior odd sepal prolonged into
a long spur; quincuncial or imbricate; blue or violet. Corolla: 4 petals, gamopetalous, valvate; two
posterio-lateral petals fuse to form the spur, and this spur is situated inside the spur formed by the
posterior sepal; sometimes only 2 petals. Androecium: Stamens numerous, usually 15, in 5 groups of
3 each, polyandrous; anthers adnate or basifixed, dithecous, extrorse. Gynoecium: Monocarpellary,
superior, unilocular, with many ovules, marginal placentation; style reduced; stigma simple. Fruit:
Follicle.
Floral Formula: Br, Brl, , , K5, C(4) or (2), A15 or μ, G1.
21.7 NYMPHAEACEAE WATER LILY FAMILY


Aquatic perennial herbs; leaves with long petiole, peltate or pseudopeltate, with latex; petals, stamens,
and sometimes sepals numerous; marginal or parietal placentation.

Nymphaea, Nelumbo, Euryale and Nuphar.
21.7.4 Size, Distribution and General Information

This family of subclass Polypetalae is represented by only 8 genera and over 90 species. Members are
cosmopolitan in distribution, except in very cold regions, and are favourite plants for garden pools.
Flowers of water lily (Nymphaea) are variously coloured and open just above the water surface in
ponds while that of crow lily (Nuphar) are golden yellow and extend up and out of the water in the
pond. Nelumbo and Nymphaea lotus are the “sacred lotus” plants of the family. Indian lotus, found
commonly in shallow lakes and ponds, is Nelumbo nucifera. Nymphaea nouchali is the Indian red
water-lily. Victoria is a native of South America.

General Habit Aquatic, freshwater, perennial herbs with large rhizomes; sometimes annual (Euryale);
submerged or free-floating in ponds, shallow lakes, etc.
Root Adventitious, branched clusters of fibrous roots develop from the root stocks or rhizomes.
Stem Perennial, branched, rhizomatous, creeping in Nymphaea but somewhat erect in Victoria and
Brasenia; full of aerenchyma, laticiferous vessels and intercellular spaces.
Leaf Simple, alternate, peltate or cordate, long petioled; submerged, floating or emersed (Nelumbo);
large, reaching up to 2 metres in Victoria; smooth and waxy but lower portion prickly (Euryale); in
Nymphaea, the mature leaves may be orbicular, sagittate or oval; Cabomba shows heterophilly.
228 Plant Taxonomy
Inflorescence Solitary and axillary.

Flower Solitary, bisexual, actinomorphic; large, showy, long-
peduncled; usually hypogynous but epigynous in Victoria and
perigynous in Nymphaea; often calyx and corolla are poorly dif-
ferentiated and are represented by perianth of 6 to numerous free
tepals.
Calyx 3 to many sepals, polysepalous; in Nuphar sepals are yel-
low and larger than petals; sepals 3 in Brasenia, 4 in Nelumbo
and Nymphaea (Fig. 21.8) and 5 in Barclaya.
Corolla 3 to many petals, polypetalous; petals frequently grading
Fig. 21.8 Floral diagram of into the stamens and therefore inner petals often form petaloid
Nymphaea nouchali. staminodia; white or variously coloured and showy; a nectary is
present on abaxial side in Nuphar; imbricate.
Androecium 3 to many, distinct stamens; filaments usually extend as sterile appendage beyond the
anther sacs; anthers dithecous, basifixed or adnate, introrse; dehiscence longitudinal; in Victoria the
inner stamens are sterile; in Nelumbo and Nymphaea the stamens are spirally arranged.
Gynoecium Carpels 2 or 3 (Cabomba) to numerous (Nelumbo, Nymphaea); each carpel with 1 loc-
ule, numerous ovules, marginal or parietal placentation; ovary superior but inferior in Victoria and
Euryale; stigmas mostly 1 and discoid or as many as 35 and radiate; numerous carpels of Nymphaea
and Nelumbo remain embedded separately on upper flat surface of swollen receptacle.
Fruit and Seed Fruit a spongy berry (Nymphaea) or follicle (Cabomba); seeds 1 to many per locule;
with straight embryo and starchy endosperm; seeds sometimes arillate (Nymphaea).
Pollination and Dispersal Large, too bright, showy and often scented flowers of this family show
entomophily, i.e. pollinated by insects such as beetles, small flies, etc. Some Nymphaea species are
self-pollinated while Nuphar, Victoria and remaining species of Nymphaea are cross-pollinated.
Dispersal in this family mainly takes place by water. However, some species of Cabomba and
Nuphar are dispersed by ducks.
General Floral Formula ≈, , K3 –μ, C3 –μ, A3 –μ, G3–μ or (3 – μ).

• Ornamentals: Because of their beautiful, showy or sweet-scented flowers, several
Nymphaeaceae members are of great ornamental value for aquatic gardening. Chief among
them are sacred ‘lotus of India’ or ‘Kamal’ (Nelumbo nucifera), Indian ‘red water-lily’
(Nymphaea nouchali), Indian ‘blue water-lily’ (Nymphaea stellata), ‘Pandharen-Kamal’
(Nymphaea alba) and ‘Royal water-lily’ (Victoria regia). Nelumbo lutea has yellow flow-
ers while Brasenia schreberi has purple flowers and Nymphaea rubra has red flowers.
Nymphaea coerulea with blue flowers, N. capensis with sky-blue flowers, N. odorata with
pink flowers and N. mexicana with bright yellow flowers are other species of ornamental
value.
• Edible Products: (i) Seeds of ‘gorgan nut’ or ‘Makhana’ (Euryale ferox) are roasted and
eaten; doctors recommend them for invalids. (ii) The vegetable ‘Kamal-Kakri’ is actually
the rhizome of Nymphaea nouchali. (iii) Fruiting torus, flowers, rhizomes, young leaves and
petioles of Nelumbo nucifera are used as vegetable. (iv) Rhizomes of Nymphaea stellata are
also edible. (v) Seeds of Victoria regia are also roasted and eaten.
• Medicinal Value: (i) Flowers of Nelumbo nucifera are recommended as a cardiac tonic
and also used in curing diseases of liver, and skin and its dried and powdered rhizome to
cure piles. (ii) Rhizomes of Nymphaea nouchali are used to cure dysentery and diarrhoea.
(iii) Flowers of Nelumbo nucifera yield a valuable perfume. (iv) Nymphaea stellata is used
to relieve cardiac pains.

On the basis of the floral construction, family Nymphaeaceae is divided into three subfamilies, i.e.
Cabomboideae, Nelumboideae and Nymphaeoideae. Apocarpous condition is found in Cabomboideae
and Nelumboideae while syncarpous condition is observed in Nymphaeoideae. Cabomboideae have
cyclic trimerous flowers whereas Nelumboideae have spiral pleiomerous flowers. All these three
subfamilies have been treated as independent families by Bessey (1915) under the names Cabombaceae
and Nelumbaceae belonging to order Ranales, and Nymphaeaceae belonging to order Rhoeadales.
Several taxonomists treated them on the line suggested by Bessey.
Cronquist (1981) included Nymphaeaceae under order Nymphaeales along with four more fami-
lies (Nelumbonaceae, Barclayaceae, Cabombaceae and Ceratophyllaceae). However, Thorne (1983)
treated Nelumbonaceae under order Nelumbonales while Nymphaeaceae and Cabombaceae under
order Nymphaeales.
Bentham and Hooker treated Nymphaeaceae under order Ranales, and subfamily Cabomboideae
forms a link between Nymphaeaceae and Ranunculaceae. Presence of trimerous flowers and scattered
vascular bundles in Cabomboideae also suggest its affinity with monocotyledons. Nymphaeaceae
show affinity with Berberidaceae in possessing scattered vascular bundles and arillate seeds, and
also with Papaveraceae in having latex, large and discoid stigma and type of placentation:
Nymphaeales have been treated as probable ancestors of monocotyledons by Cronquist (1967)
because of their aquatic occurrence, lack of vessels, absence of cambium, presence of uniaperturate
pollen and laminar placentation.
21.7.8 Description of a Common Plant

1. Nelumbo nucifera Gaertn. (‘Sacred lotus’ or ‘Kamal’; Fig. 21.9)
Habit: Aquatic, perennial herb. Stem: Branched, rhizomatous. Leaves: Simple, large, forming a
triad, of which one is present on the lower side of rhizome and scaly, another is present on the
upper side of rhizome, and third leaf is large and reaches up to the water surface; latter is long-
petioled, stipulate, peltate; upper surface waxy; multicostate reticulate. Inflorescence: Solitary axil-
lary. Flower: Ebracteate, pedicellate, pedicel long; large, complete, hermaphrodite, actinomorphic,
hypogynous, showy. Calyx: 4 sepals, free, petaloid or greenish pink, imbricate. Corolla: Numerous
petals, polypetalous, spirally arranged, pink. Androecium: Numerous stamens, polyandrous, spirally
230 Plant Taxonomy
leaf flower bud
stamen
thalamus
carpels A Petal
(gradually changing
into stamen)
Fruit
Flower Bud
A Flower
anthers
appendage
thalamus
stigma
carpel
Floral Diagram L.S. Thalamus A Flower A Stamen

(petals removed)
Fig. 21.9 Nelumbo nucifera Gaertn.
arranged; filaments long, anthers basifixed, connective is prolonged as an appendage. Gynoecium:

Polycarpellary, apocarpous, superior; carpels embedded in the upper part of an obconically flat but
spongy receptacle; unilocular, one ovule in each locule; stigma flat.
Floral Formula: Ebr, ≈, , K4, C μ, A μ, G μ.
21.8 PARIETALES
1. Pentamerous flowers with their calyx showing imbricate aestivation.
2. Stamens as many as petals or more
3. Ovary tricarpellary, syncarpous, unilocular
4. Numerous ovules showing parietal placentation
5. Seeds endospermic
Bentham and Hooker divided cohort Parietales into 9 orders (=families), namely Cruciferae,
Papaveraceae, Violaceae, Bixineae, Cannellaceae, Sarraceniaceae, Capparidaceae, Resedaceae and
Cistineae. Engler and Diels, however, divided order Parietales into 10 suborders composed of 31
families. Lawrence (1951), while following Engler and Diels, however, commented that “available
evidence indicates that the order is not a phylogenetic taxon, and the realignment of the families
into several orders is to be expected. In some instances this may result in transfer of families to
existing orders and in others in the establishment of new orders”.
Only Papaperaceae, Fumariaceae, Capparidaceae, Cruciferae (Brassicaceae) and Violaceae are
treated in this text.
21.9 PAPAVERACEAE POPPY FAMILY

Polypetalae, Thalamiflorae, Parietales.

Herbs; leaves alternate, often lobed or divided, exstipulate; flowers bisexual; poisonous latex usually
present; petals rolled or crumpled in the bud; stamens numerous; ovary unilocular; fruit capsule.

Argemone, Eschscholzia, Papaver.

Poppy family is represented by about 26 genera and 280 species (Hickey and King, 1988) distributed
chiefly in Northern Hemisphere. Western North America and eastern Asia are the main centres of
its distribution. Representatives of this family also occur in Europe, South Africa, Australia and
even in arctic regions.
Only 5 genera and about 20 species have so far been reported from India. Meconopsis with
about 26 species grows wild in some parts of Himalayas. Argemone mexicana occurs wild in waste
fields while Papaver somniferum (opium poppy) is cultivated widely in India. The drug “opium” is
obtained by cutting notches in the half-ripened capsules, from which the latex exudes.
232 Plant Taxonomy

General Habit Mostly annual or perennial herbs with milky or coloured latex; rarely woody shrubs
(Dendromecon) or tree-like, reaching up to 10 metre high (Bocconia arborea).
Roots Extremely branched, usually surface feeder.
Stem Herbaceous, rarely woody; branched, erect, cylindrical; glabrous or sometimes hairy.
Leaves Alternate (rarely the upper ones whorled), basal or cauline; usually much divided or lobed
and pinnately cut; exstipulate; with milky or coloured sap or latex; the latex is white in Papaver,
yellow in Argemone, orange in Chelidonium, watery in Eschscholzia, and bright orange-red in
Sanguinaria.
Inflorescence Solitary showy flowers in most of the species; racemes or cymes in Meconopsis;
racemose panicles in Bocconia; umbellate clusters in Chelidonium.
Flower Ebracteate, pedicellate, bisexual, actinomorphic,
complete, hermaphrodite, bi- or trimerous; large and showy;
hypogynous but perigynous in Eschscholzia.
Calyx Sepals 2 to 3, polysepalous but connate in Eschscholzia;
caducous i.e. fall off very soon; imbricate.
Corolla Petals 4 to 6 or more, polypetalous, often crumpled
(full of folds) in the bud (Fig. 21.10); in one or two or rarely
more whorls; imbricate; bright coloured; petals absent in
Bocconia and Macleaya.
Androecium Stamens numerous, free, arranged in closed
spiral; filaments short; anthers elongate, basifixed, extrorse;
dehiscence longitudinal.
Gynoecium A compound pistil of 2 or more fused carpels,
ovary superior, with one locule containing numerous ovules,
Fig. 21.10 Floral diagram of parietal placentation; ovary half-inferior in Eschscholzia;
Papaver rhoeas. ovary rarely becomes multilocular by fusion of intruding
placentae; style reduced; stigmas as many as carpels; ovules
anatropous or slightly campylotropous; rarely only one basal ovule in Bocconia.
Fruit and Seed Fruit a capsule, opening by valves (Argemone) or pores (Papaver); seeds numerous,
with copious and oily endosperm and small embryo.
Pollination and Dispersal Pollination is entomophilous. Flowers are mostly large and conspicuous,
but usually do not contain any nectar, and are visited mainly by pollen-seeking insects. In Papaver,
the flowers are homogamous and both self- and cross-pollination take place. Dispersal of seeds is
affected usually by wind or by animals, birds or human beings.
General Floral Formula Ebr, ≈, , K2 – 3 or (2), C2+ 2 or 3 + 3, A μ, G(2 – μ).
• Opium poppy (Papaver somniferum), cultivated in the warmer countries for the universally
known drug opium, is the most important plant of this family. Opium is obtained from the
milky latex of unripe capsules of this plant. Morphine content ranges from 5–6% in unripe
fruits, while only 0.8–1% in ripe fruits. Morphine, codeine, narcotine and papaverine are
some of the alkaloids present in this milky latex. Opium is known for its sedative properties
and its high doses first produce sleep and then depression, paralysis, unconsciousness and
ultimately death.
• The seeds of Papaver somniferum contain no opium, and are used in baking and sprinkled
on bread.
• An edible oil, obtained from the seeds of P. somniferum, is used in the preparation of animal
and human foods, and also in the manufacture of soaps, paints and varnishes.
• Wrapping paper is prepared from the pulp obtained from opium poppy.
• Family is known for its popular garden plants, such as Eschscholzia californica (Californian
Poppy), Argemone mexicana (Prickly poppy), Macleaya cordata (Plume poppy), Meconopsis
(Blue poppy), Dendromecon rigida (Bush poppy), Sanguinaria canadensis (Blood root) and
Chelidonium (Celandine).
• True poppies are species of Papaver, such as P. nudicaule (Iceland poppy, orange red flow-
ers), P. orientale (Oriental poppy, bright red flowers), P. rhoeas (Corn poppy, red or deep
purple flowers), etc.
• A non-edible oil, obtained from the seeds of Argemone mexicana, is used as an illuminant
and in skin infections. It is also mixed to adulterate mustard oil.
• An emetic drug, ‘Sanguinarine’, is obtained from the rhizome and roots of Sanguinaria
canadensis. It is used in dyspepsia i.e. indigestion.

Bentham and Hooker included Papaveraceae under order Parietales whereas Engler and Prantl (1887–
1899) treated it under order Rhoeadales. Takhtajan (1980) also treated it under Rhoeadales whereas
Cronquist (1981) included it under order Papaverales along with one more family Fumariaceae. Thorne
(1983) treated it under suborder Papaverineae of order Berberidales.
Family Papaveraceae shows most close affinities with Ranunculaceae, mainly in having hemicy-
clic flowers, numerous stamens with extrorse anthers, numerous carpels, and the seeds with copious
endosperm and very small embryo. Papaveraceae also shows affinities with Brassicaceae (Cruciferae)
in possessing syncarpous, unilocular ovary and parietal placentation. Flowers of Platystemon establish
a link between Papaveraceae and Nymphaeaceae.

1. Argemone mexicana L. (Prickly Poppy or Peeli Kataili or Jangli Posth; Fig. 21.11)
Habit: A spiny or prickly annual herb. Stem: Herbaceous, erect, branched, cylindrical, solid, green,
spiny, with yellow latex. Leaf: Simple, alternate, exstipulate, sessile, semi-amplexicaul; margin
234 Plant Taxonomy
stigma
petal stigma
petal
stamen
spinous ovary
sepal ovary stamen
pedicel
L.S. Bud pedicel
ovules ovary
wall
L.S. Flower
anther lobe
connective
ovary ovary
filament
thalamus
pedicel A Stamen
Gynoecium
T.S. Ovary
flower
leaf
stem
Fig. 21.11 Argemone mexicana L.
dissected and spinous; apex acute; surface spiny; unicostate reticulate. Inflorescence: Solitary ter-
minal or solitary axillary. Flower: Ebracteate, pedicellate, complete, hermaphrodite, actinomorphic,
trimerous, hypogynous, yellow. Calyx: Sepals 3, polysepalous, imbricate or twisted; spiny; caducous,
contains a horn-like spine at the apex. Corolla: Petals 6, polypetalous, arranged in two whorls of
three each, caducous, crumpled in the bud, yellow. Androecium: Stamens numerous, polyandrous,
filaments long, dithecous, basifixed, extrorse. Gynoecium: 4 to 6 carpels, syncarpous, superior, uni-
locular, many ovules, parietal placentation; style reduced; stigma 4–6 lobed and hood like; ovary
covered with prickles. Fruit: Capsule. Seed: Black, small, endospermic.
Floral Formula: Ebr, ≈, , K3, C3 + 3, A μ, G (4 – 6).
2. Eschscholzia californica Chamb. (Californian Poppy; Fig. 21.12)
Habit: Herb. Root: Branched, tap root. Stem: Erect, herbaceous, branched, green, solid, glabrous.
Leaf: Cauline and ramal, simple, alternate, exstipulate, very much dissected or decompound, leaf
base sheathing; acute, glabrous; unicostate reticulate. Inflorescence: Solitary terminal or solitary axil-
lary. Flower: Ebracteate, pedicellate, complete, hermaphrodite, actinomorphic, bimerous, perigynous,
ovary wall
petal placenta
anther locule
ovules
stigma stigma
style
T.S. Ovary
ovary style
ovule
thalamus ovary
pedicel
L.S. Flower flower
Gynoecium
leaf
stem
Flowering Branch
Floral Diagram
Fig. 21.12 Eschscholzia californica Chamb.

236 Plant Taxonomy
cyclic, yellow. Calyx: Sepals 2, raised on cup-like projection of floral axis; fused to form a pointed
hood, specially in bud condition; caducous. Corolla: Petals 4, polypetalous, arranged in 2 whorls
of 2 each, imbricate; bright yellow. Androecium: Stamens numerous, polyandrous, arranged at the
brim of the cup; filaments short, dithecous, basifixed, extrorse. Gynoecium: Bicarpellary, syncarpous,
half-inferior, unilocular, ovules numerous; parietal placentation; style short; stigma 2, each is bifid;
out of 2 portions of stigma one is longer than the other. Fruit: Capsule.
Floral Formula: Ebr, ≈, , K(2), C2 + 2, A μ, G(2)–.
21.10 FUMARIACEAE FUMARIA FAMILY OR FUMITORY FAMILY

Treated as a tribe of family Papaveraceae by Bentham and Hooker.

Herbs with watery juice but latex absent; leaves usually variously divided; flowers zygomorphic;
petals spurred; stamens 6, in 2 groups; gynoecium bicarpellary.

Fumaria, Corydalis, Dicentra, Hypecoum.

Fumariaceae, often treated as a sub-family (Fumarioideae) of Papaveraceae, is represented by
about 19 genera and over 400 species. It is distributed in temperate North America, Asia, Africa
and Europe. 4 genera and over 40 species of this family have been reported from India. The three
best known genera with their reported number of species are Corydalis (300), Fumaria (50) and
Dicentra (22).

General Habit Annual (Fumaria indica) or perennial herbs, with watery juice; sometimes climbers
(Corydalis, Fumaria officinalis); instead of latex vessels of Papaveraceae, oil-containing sacs are
present in Fumariaceae.
Root Branched tap roots, sometimes tuberous (Corydalis).
Stem Herbaceous; erect or climbing by leaf or petiole tendrils; sometimes tuberous (Corydalis) or
bulbous (Dicentra); watery juice present.
Leaves Alternate, rarely opposite; simple with much divided blades (Fumaria indica); petiolate;
sometimes with tendril-like leaf axes; unicostate reticulate.
Inflorescence Usually racemose racemes; rarely subumbellate (Corydalis)
Flower Bracteate, shortly pedicellate, complete, hermaphrodite; laterally or transversely zygomor-
phic; dimerous; hypogynous; cyclic.
Calyx Sepals 2, small, free; medianly placed; caducous or deciduous.

Corolla Petals 4, polypetalous, in 2 whorls of 2 each; one (Fumaria, Corydalis) or both (Dicentra)
outer petals more or less spurred at base (Fig. 21.13A, B); two inner petals are smaller and often
fused around the anthers; in Hypecoum the outer petals are trilobed at the apex while the inner
petals are tripartite (Fig. 21.14).
Androecium Androecium consists of 2 groups or bundles, located opposite to the inner petals; each
group divided into 3 parts at the apex, i.e. central part bearing a complete or dithecous anther and
the 2 lateral parts, each bearing monothecous or half an anther (Fig. 21.15); stamens are 4 and anti-
petalous in Hypecoum; polyandrous; extrorse; a basal nectary is attached with either one (Fumaria,
Corydalis) or both (Dicentra) the groups of stamens.
Some views regarding the morphological nature of androecium in Fumariaceae are
undermentioned:
1. Asa Gray believed that each lateral group or bundle represents a single stamen which has
become tripartite.
2. De Candolle opined that each lateral group of stamens consists of one lateral stamen and 2
half-stamens derived from 2 posterior and 2 anterior stamens of a 4-stamened androecium,
as of Hypecoum.
3. Arber considered them as 6 stamens, of which 2 are normal with dithecous anthers and 4
are reduced with monothecous anthers.
4. Eichler has been of the view that Fumariaceae members have only 2 dithecous stamens, and
the 4 monothecous parts attached with them are their stipular appendages.
5. Norris believed that each lateral group of stamens is a double structure, made up of 2 stamens
with monothecous anthers. It is so because each group or bundle receives 2 bundles.
Gynoecium Bicarpellary, syncarpous, superior, unilocular, one to many ovules in locule, parietal
placentation; style filiform; stigma capitate or slightly bilobed.
Fruits and Seeds Fruit a slender capsule or one-seeded nutlet; seeds black and shiny, often with an
aril or caruncle, each contains a minute embryo and fleshy endosperm.
A B
Fig. 21.13 Floral diagrams— Fig. 21.14 Floral diagram of Hypecoum.

A: Corydalis; B: Dicentra.
238 Plant Taxonomy
Pollination and Dispersal Nectary, attached with one (Fumaria, Corydalis) or both (Dicentra) the
lateral groups of stamens, secretes honey which attracts the nectar-seeking insects helpful in pol-
lination. Dispersal of seeds takes place by ants, birds or human being.
General Floral Formula Br, , , K2, C2 + 2 or 2 + (2), A(3) + (3), G (2).
• Alkaloid protopine, found in Hypecoum procumbens and H. leptocarpum, is used in high

blood pressure.
• Some species of Corydalis and Dicentra are grown as ornamental plants.
• Corydalis racemosa is used in eye diseases while C. govaniana roots are used against
syphilis.
• Dried plants and seeds of Fumaria indica are used to cure fever and as blood purifier.

Botanists, such as Bentham and Hooker (1862–1883), Rendle (1925), Engler and Prantl (1931),
Melchior (1964), etc. treated this family as a tribe or subfamily of family Papaveraceae. However,
Hutchinson (1959) provided it the status of an independent family (Fumariaceae) under order
Rhoeadales. Hutchinson’s treatment is followed by most of the recent botanists, including Cronquist
(1981) who discussed Fumariaceae and Papaveraceae as two independent families of order Papaverales.
Hypecoum (Fig. 21.14) forms a connecting link between Fumariaceae and Papaveraceae. Moreover,
Fumariaceae differ from Papaveraceae in showing absence of latex, and presence of variously divided
leaves, zygomorphic flowers, spurred petals and stamens in two groups.
21.10.8 Description of Common Plant

1. Fumaria indica L. (= F. parviflora; Pit-papra, Fig. 21.15)
Habit: Annual herb. Root: Well-branched tap root. Stem: Herbaceous, branched, green, without latex.
Leaf: Alternate, exstipulate, simple or decompound, petiolate, finely-dissected into several lobes or
segments; each lobe entire, acute, unicostate reticulate. Inflorescence: Racemose raceme. Flower:
Bracteate, pedicellate, complete, zygomorphic, dimerous, hypogynous, white or purple-pink. Calyx:
Sepals 2, free, membranous, anterio-posterior, caducous, valvate. Corolla: Petals 4, arranged in 2
whorls of 2 each; one of the outer lateral petals is spurred; inner whorl petals are smaller and placed
anterio-posterior. Androecium: Stamens in 2 groups; each group contains 1/2 + 1 + 1/2 stamens;
filament of the stamen opposite to the spurred petal has a yellow-green nectary; basifixed, extrorse.
Gynoecium: Bicarpellary, syncarpous, superior, unilocular, many ovules, parietal placentation, stigma
lobed. Fruit: Nut.
Floral Formula: Br, , , K2, C2 + 2, A 12 + 1 + 12 + 12 + 1 + 12 , G (2).
ovule
ovary
wall
leaf
T.S. Ovary
flowers
stigma
style
stamen
ovary
spur
nectary stem
L.S. Flower Flowering Branch
inner whorl
of petals
outer petal
stigma outer
petal
style
ovary pedicel
nectary
Floral Diagram Parts of a Flower
Fig. 21.15 Fumaria indica L.
21.11 CAPPARIDACEAE THE CAPERS FAMILY

240 Plant Taxonomy

Herbs, shrubs or climbers; leaves alternate, simple or compound; flowers usually actinomorphic;
presence of androphore and gynophore; fruit capsule or berry.

Cleome, Capparis, Crataeva.

42 genera and over 900 species of this family are distributed mostly in the tropics and warm temper-
ate regions of the world. Only 7 genera and over 65 species have so far been reported from India,
mostly from western and southern parts. Quite a large number of plants occur in xeric conditions.
Over 35 species of Capparis have been reported from India. Roydsia is very common in Assam.

General Habit Herbs (Cleome viscosa, C. gynandra), shrubs (Capparis), lianas (Maerua are-
naria), small tree (Niebuhria linearis) or tall tree (Crataeva religiosa). Mostly grow wild and are
xerophytic.
Root Well-branched, deep feeder, tap root.
Stem Aerial, erect, herbaceous or woody (Capparis), rarely climbing (Maerua arenaria), hairy or
covered with thorns (Capparis aphylla).
Leaf Alternate, simple (Capparis zeylanica, Maerua) or palmately compound (Cleome, Crataeva);
usually stipulate; stipules may be foliaceous (Cleome) spinose (Capparis) or setaceous (Niebuhria);
in Capparis decidua leaves are reduced, scaly and caducous.
Inflorescence Generally solitary or in racemes (Cleome, Polanisia), rarely shortly peduncled simple
umbel (Capparis sepiaria).
Flower Bracteate, ebracteolate, complete, hermaphrodite, actinomor-
phic, tetramerous, hypogynous; in some (Pteropetalum, Emblingia)
flowers are medianly zygomorphic; incomplete due to the absense of
petals in Roydsia and Niebuhria; a disc is usually present; thalamus
of several genera is elongated into androphore and gynophore, col-
lectively called androgynophore.
Calyx 4, free or rarely basally fused sepals, arranged in 2 whorls
of 2 each; in Capparis posterior sepal forms a hood-like structure;
valvate or imbricate; sepals are 5 and basally fused in Emblingia;
in Roydsia 6 sepals are arranged in 2 whorls of 3 each.
Corolla 4, free, diagonally placed petals; valvate (Cleome) or imbri-
cate (Capparis, Fig. 21.16); petals are 2 in Cadaba and Emblingia
and none in Roydsia; in Pteropetalum, the 2 posterior petals are
Fig. 21.16 Floral diagram larger than the 2 anterior ones.
of Capparis.
Androecium Stamens 4 (Cleome tetrandra) to numerous (Crataeva, Capparis), polyandrous; anthers

dithecous, introrse; in some genera (Cleome gynandra) the receptacle of the flower between corolla
and stamens gets elongated in the form of androphore; stamens are 6 in Cleome spinosa but never
tetradynamous.
Gynoecium Usually bicarpellary, rarely tetra- to multicarpellary; syncarpous; in some genera the
receptacle of the flower gets elongated between androecium and gynoecium in the form of gynophore;
superior; unilocular, rarely falsely bilocular; a few to many ovules, parietal placentation; style short;
stigma capitate or 2-lobed.
Fruit and Seed Fruit berry (Capparis, Crataeva), capsule (Cleome, Polanisia), drupe (Roydsia) or
nut (Emblingia). Seeds non-endospermic, kidney-shaped, with folded embryo as in Brassicaceae.
Pollination Entomophilous.
General Floral Formula Br, Ebrl, ≈, , K2 + 2, C4, A4– μ, G(2 – 4).

Members of Capparidaceae are of little economic importance.
• Ornamentals: Capparis sepiaria, C. spinosa, Cleome chelidonii, C. spinosa (Capers), and
Roydsia suaveolens are grown as ornamental plants.
• Medicinal: (i) Capparis zeylanica leaves are used against boils and stomach ache; (ii)
Cleome gynandra (syn. Gynandropsis pentaphylla) leaves and seeds are used for expelling
roundworms, and juice from its roasted leaves is used to cure earache; (iii) Crataeva religiosa
bark is used for increasing appetite and reduction of bile secretion; (iv) Maerua arenaria
roots are used as a tonic and stimulant.
• Edibles: (i) Capparis spinosa fruits are picked and form the ‘Capers of Commerce’; (ii)
Unripe fruits of Capparis aphylla (syn. C. decidua) and C. zeylanica are also pickled and
eaten in India; (iii) Seeds of Cleome viscosa (‘Hurhur’) are used in curries; (iv) Unripe
fruits of Maerua arenaria are boiled and eaten.

Because of the presence of parietal placentation and commissural stigmatic lobes, Capparidaceae
are related closely to the Parietalean families, such as Brassicaceae, Moringaceae, Papaveraceae,
Resedaceae, etc.
Bentham and Hooker treated Capparidaceae under order Parietales while Engler and Prantl
(1931) placed them under order Rhoeadales. Capparidaceae have been treated under a new name
Capparaceae belonging to order Capparales by majority of the recent taxonomists, including Takhtajan
(1969), Cronquist (1981) and Thorne (1983). Capparidaceae are usually divided into two tribes i.e.
Cleomeae, including herbaceous plants and, Cappareae including woody plants.
242 Plant Taxonomy
stigma ovary wall

style locule
ovule
ovary
anther lobe glands
ovule
filament
gynophore stigma
style T.S. Ovary
petal
sepal ovary
androphore glands
pedicel
L.S. Flower (diagrammatic) Gynoecium

flower
fruit
leaflet stem
Floral Diagram
Flowering Branch
Fig. 21.17 Cleome gynandra D.C.

1. Cleome gynandra D.C. (syn. Gynandropsis pentaphylla L.; Fig. 21.17)
Habit: Annual, erect herb. Stem: Herbaceous, erect, branched hairy. Leaf: Alternate, exstipulate
petiolate, palmately compound; leaflets sessile, ovate to obovate, acute, hairy, unicostate reticulate.
Inflorescence: Corymbose raceme. Flower: Ebracteate, pedicellate, complete, hermaphrodite, acti-
nomorphic, hypogynous, tetramerous; androphore and gynophore present. Calyx: 4 sepals, free,
arranged in 2 whorls of 2 each, imbricate. Corolla: 4 petals, free, valvate, clawed, white. Androecium:
6 stamens, polyandrous, dithecous, dorsifixed, introrse; floral axis is elongated between corolla and
stamens in the form of androphore; filaments long. Gynoecium: Bicarpellary, syncarpous, superior,
unilocular, many ovules, parietal placentation; floral axis is elongated between androecium and ovary
in the form of gynophore; style reduced; stigma capitate. Fruit: Capsule.
Floral Formula: Ebr, ≈, , K2 + 2, C4, A6, G(2).
21.12 BRASSICACEAE OR CRUCIFERAE MUSTARD FAMILY


Herbs with a smelling watery juice; flowers of 4 sepals, 4 petals and 6 stamens; corolla cruciform;
stamens tetradynamous; ovary with a false septum; fruit siliqua or silicula.

Alyssum, Brassica, Capsella, Coronopus, Eruca, Iberis, Raphanus, Sisymbrium.

This cosmopolitan family of about 375 genera and over 3200 species, occur chiefly in north tem-
perate regions, specially in the Mediterranean region. About 50 genera and over 140 species have
so far been reported from India. Some of the larger genera, along with their reported number of
species from the world (Hickey and King, 1988) in parenthesis, include Draba (300), Cardamine
(l60), Lepidium (150), Alyssum (150), Arabis (120), Erysimum (100), Sisymbrium (90), Heliophila
(75), Rorippa (70), Brassica (50), Iberis (30) and Raphanus (8).

General Habit Annual, biennial, or perennial herbs with a pungent watery sap; herbage commonly
covered with branched or stellate unicellular hairs; rarely subshrubs.
Root Tap root which in some genera gets swollen due to stored food and becomes fusiform (rad-
ish) or napiform (turnip).
Leaf Mostly simple, alternate, often dissected, exstipulate; rarely opposite or subopposite; pubes-
cence of simple to stellate hairs; radical leaves form a rosette whereas cauline leaves are alternate
or opposite.
Inflorescence Typically a raceme or corymb (Iberis) and nearly always without bracts or
bracteoles.
Flower Ebracteate, ebracteolate, complete, bisexual, actinomorphic, rarely zygomorphic (Iberis,
Fig. 21.18A); tetramerous; hypogynous.
Calyx 4 sepals, polysepalous, in 2 whorls of 2 each, outer whorl is anterio-posterior.
Corolla 4 petals, polypetalous, cruciform (arranged in a cross), alternate with sepals, often clawed;
2 anterior petals in Iberis are larger (Fig. 21.18A); petals reduced or scale-like in Coronopus
(Fig. 21.18B); petals completely absent in Lepidium.
Androecium 6 stamens, arranged in 2 whorls; an outer whorl of 2 short stamens, and an inner
whorl of 4 long stamens (i.e. tetradynamous); anthers dithecous (rarely monothecous); introrse,
244 Plant Taxonomy
A B
Fig. 21.18 Floral diagrams—A: Iberis amara and, B: Coronopus didymus syn. Senebiera didyma.
longitudinally dehiscent; stamens are only 4 in some species of Nasturtium while only 2 in Coronopus
didymus (Fig. 21.18B) and Lepidium and up to 16 in Megacarpaea polyandra.
Gynoecium Bicarpellary, syncarpous, superior; unilocular but becoming bilocular due to the develop-
ment of a false septum or replum; replum unites the two parietal placentae; numerous anatropous or
campylotropous ovules; parietal placentation; styles 1 or obsolete; stigma capitate to bilobed; carpels
3 in Lepidium sativum and 4 in Tetrapoma.
Fruit and Seed Fruit siliqua (Arabis, Brassica, Cheiranthus cheiri) or silicula (Capsella bursa-
pastoris, Lunaria), or occasionally a nut (Crambe) or rarely achene-like (Isatis tinctoria). Seeds
are small, non-endospermic, with curved embryo and often mucilaginous testa. Fruits and seeds are
characters of systematic importance in this family.
Pollination and Dispersal Pollination is entomophilous, and dispersal of seeds may be by birds,
wind, water or cattles.
General Floral Formula Ebr, Ebrl, ≈, , K2 + 2, C4, A2+ 4, G(2)

Brassicaceae are of considerable economic importance for providing us a variety of vegetables,
oils, food crops, ornamentals, and weeds.
1. Vegetables: Several common vegetables of daily use belong to this family. Some of them
are Mooli (Raphanus sativus, radish), Shaljam (Brassica rapa, turnip), Phool Gobhi (B.
oleracea var. botrytis, cauliflower), Band Gobhi (B. oleracea var. capitata, cabbage), Ganth
Gobhi (B. oleracea var. gongylodes, Knol-knol), Button Gobhi (B. oleracea var. gemmifera,
Brussel’s sprouts), Kadamsag (B. oleracea var. acephala), Sarson (B. campestris var. sarson,
mustard) and Sengri (B. sativus var. caudatus).
2. Oils: Fatty oil, used for cooking, pickles, burning, massaging, etc. is obtained from the
seeds of several species of this family. Mustard oil is obtained from several species of
Brassica, such as Brassica campestris var. sarson (mustard or yellow sarson) and B. camp-
estris var. toria (Toria or Indian rape). Oil is also obtained from the seeds of B. juncea
var. cuneifolia (Rai), B. nigra (Kali Rai or black mustard), B. alba (white or Safed sarson)
and Eruca sativa (Taramira). The oil cake left after the extraction of oil, is a good cattle
feed.
3. Ornamentals: Among the well-known ornamentals are candytuft (Iberis), honest (Lunaria),
sweet alyssum (Lobularia), wall flower (Cheiranthus), stocks (Mathiola), etc.
4. Medicinal: Majority of Crucifers produce an abundance of vitamin C. Characteristic pun-
gent odor of its members is because of sulphur compounds. Cauliflower and a few other
Crucifers have recently attracted botanists as possible anticancer food items.
5. Weeds: Troublesome weeds of cultivated fields and lawns include Brassica arvensis,
Capsella bursa-pastoris (shepherd’s purse), Barbarea, Lepidium virginicum (pepper grass),
Coronopus didymus and Sisymbrium.

Cruciferae has been placed in order Parietales by Bentham and Hooker while Engler and Prantl placed
it under order Rhoeadales. Hutchinson (1959) placed it under order Cruciales while Cronquist (1981)
and Thorne (1983) placed it under order Capparales. Some taxonomists believe that Cruciferae has
been derived from Papaveraceae whereas others believe it to be derived from Capparidaceae.

Brassica campestris var. sarson (Sarson; Fig. 21.19)
Stem: Herbaceous, erect, branched, smooth. Leaf: Alternate, exstipulate, simple, sessile or subsessile,
lower leaves lyrate, upper ones ovate to lanceolate; serrate, acute, unicostate reticulate. Inflorescence:
Racemose raceme. Flower: Ebracteate, ebracteolate, pedicellate, complete, actinomorphic, hermaph-
rodite, tetramerous, hypogynous, yellow. Calyx: 4 sepals, in 2 whorls of 2 each; free, slightly petaloid.
Corolla: 4 petals, polypetalous, clawed, cruciform, valvate or imbricate. Androecium: 6 stamens,
polyandrous, tetradynamous; in 2 whorls of outer 2 short stamens and inner 4 long stamens; dith-
ecous, basifixed, introrse. Gynoecium: Bicarpellary, syncarpous, superior; unilocular but development
of replum or false septum makes it bilocular; many ovules in each locule; parietal placentation; style
reduced; stigma bilobed.
Floral Formula: Ebr, ≈, , K2 + 2, C4, A2 + 4, G(2).
246 Plant Taxonomy
petal style stigma

stigma
stamen
filament
style sepal ovules ovary

petal
pedicel
A Flower nectary sepal
pedicel
L.S. Flower anther
lobe
connective
filament
A Stamen
A Mature Leaf
buds
Fruit flower
fruit
leaf
stem
Fig. 21.19 Brassica campestris L. var. sarson Prain.
21.13 VIOLACEAE VIOLET FAMILY


Herbs or shrubs; pentamerous, zygomorphic flowers; spurred corolla; 1 of the 5 stamens is frequently
spurred at the base; parietal placentation; fruit a capsule.
21.13.3 Indian Genera

Viola, Hybanthus and Vahila.

A family of about 22 genera and 900 species, Violaceae are distributed in both tropical and temper-
ate regions. Only 3 genera (Hybanthus, Vahila and Viola) have been reported from India. Famous
pansies, violas and violets belong to this family. Some of the larger genera with their approximate
number of species in parenthesis include Viola (500), Rinorea (340) and Hybanthus (150). Majority
of Indian Violaceae are found in Himalayas and hills of southern and western India.

General Habit Usually perennial or annual herbs (Viola, Hybanthus), or shrubs (Alsodeia); rarely
trees (Rinorea) or woody climbers (Agatea).
Root Branched, tap roots.
Stem Usually short, solid, erect, usually unbranched.
Leaves Alternate or rarely opposite (Hybanthus), simple; stipulate, stipules often large and leafy
(some Viola species) or small.
Inflorescence Flowers solitary or arranged in racemose racemes (Rinorea) or panicles.
Flower Bracteate, invariably bracteolate, bisexual, zygomorphic (Viola odorata, Fig. 21.20) to actino-
morphic (Rinorea); sometimes cleistogamous and hidden underground; pentamerous, hypogynous.
Calyx 5 sepals, persistent, polysepalous or slightly connate; uniformly sized; imbricate or
quincuncial.
Corolla 5 petals, mostly unequal, polypetalous; anterior petal larger and often spurred to hold the
nectar; imbricate; equal-sized petals in Rinorea and other arborescent genera.
Androecium 5 stamens, usually alternate with petals, free or connate, often forming a cylinder
round the ovary; abaxial stamens often spurred at the base; filaments very short, connective usually
prolonged at the apex, dithecous; introrse; 2 anterior stamens are spurred in Viola (Fig. 21.20).
Gynoecium Tricarpellary, syncarpous, superior, unilocular, ovules numerous, parietal placentation;
style simple; stigma capitate, truncate or lobed.
Fruits and Seeds Fruit a loculicidal capsule or rarely a berry. Seeds endospermic; smooth, winged
or with tomentum; with straight embryo; elaiosome is present in some species of Viola.
Pollination and Dispersal Pollination entomophilous in most of the Violaceae. However, cleistoga-
mous flowers (Viola bicolor) are self-pollinated. Dispersal of seeds is usually by animals, birds, or
by water.
General Floral Formula Br, Brl, or ≈, , K5, C5, A5, G(3).
248 Plant Taxonomy
petal sepal
ovules
spurred
petal
sepal
T.S. Ovary
spurred
stamen leaf Fruit
L.S. Flower
flower
spurred petal
Flower with
Sepals Removed
Floral Diagram
Flowering Plant
Fig. 21.20 Viola odorata L.

• Ornamentals: Except for garden favourites, such as pansies, violas, and violets, the family
is of little economic importance. Over 100 species of Viola are being grown as ornamental
plants. Common cultivated violet is Viola odorata and common pansy is V. tricolor.
• Medicinal: Dried flowers of Viola odorata (Vern. Banafsha) are used against cough, influ-
enza and lung troubles. Roots of Hybanthus enneaspermus are used against urinary tract
infections and gonorrhoea, and also to cure bowel troubles of children.
• Oil and Perfumery: Oil obtained from the flowers of Viola odorata are used in perfumes
and flavourings. Its leaves yield an essential oil used in scenting soaps.

Violaceae has been placed under order Parietales by Bentham and Hooker. Hutchinson (1959) placed
it under Violales and opined that Violaceae are derived from Ranalian stock through Rhoeadales.
Cronquist (1981) treated it as belonging to order Violales along with 23 more families while Thorne
(1983) discussed Violaceae under suborder Violineae of order Violales belonging to superorder
Violiflorae. Violaceae is divided into 2 subfamilies i.e. Violoideae and Leonioideae (Hickey and
King, 1988).
21.14 CARYOPHYLLINEAE
Caryophyllineae, as circumscribed by Bentham and Hooker, show following characteristics:
1. Flowers actinomorphic and bear calyx with 2–5 or rarely 6 sepals and usually as many
petals.
2. Flowers show polysepalous and polypetalous conditions.
3. Stamens typically 5 or 10, or as many as petals, or twice as many as petals, arranged in 1
or 2 whorls, obdiplostemonous.
4. Ovary unilocular; numerous ovules, free-central placentation.
5. Seeds usually with curved embryo.
Cohort Caryophyllineae includes 4 orders (=families), namely Frankeniaceae, Caryophyllaceae,
Portulacaceae and Tamariscineae. Only Caryophyllaceae and Portulacaceae are treated in the pres-
ent text. Lawrence (1951), however, treated both Caryophyllaceae and Portulacaceae under order
Centrospermae along with 8 more families, viz. Chenopodiaceae, Amaranthaceae, Nyctaginaceae,
Phytoloccaceae, Gyrostemonaceae, Achatocarpaceae, Aizoaceae and Basellaceae.
21.15 CARYOPHYLLACEAE PINK FAMILY

Polypetalae, Thalamiflorae, Caryophyllineae.

Herbs with often swollen nodes; leaves opposite decussate; inflorescence dichasial cyme or flowers
solitary; flowers obdiplostemonous; placentation free-central; fruit capsule.
250 Plant Taxonomy

Antigonon, Dianthus, Drymaria, Gypsophila, Saponaria, Silene, Spergula, Stellaria.

Represented by about 75 genera and over 2000 species (Jones and Luchsinger, 1987), Caryophyllaceae
are cosmopolitan in distribution, and mainly found in the north temperate or warm temperate regions
of the world. Over 20 genera and about 100 species have so far been reported from India. Some of
the larger genera, along with their common names and number of species in parenthesis, include
Silene (catchfly, 500), Dianthus (carnation, 300), Arenaria (sandwort, 250), Gypsophila (baby’s
breath, 125), Stellaria (chickweed, 100), Cerastium (mouse-ear chickweed, 100), Drymaria (50),
Spergularia (40), Saponaria (30), etc.

General Habit Mostly annual or perennial herbs, rarely suffrutescent shrubs; arctic forms are reduced
and possess caespitose habit.
Stem Herbaceous; rarely with a basal woody part; nodes swollen; branching usually dichotomous
but sometimes monopodial.
Leaf Simple, usually opposite decussate, rarely alternate (Dysphania), connected at the base by a
transverse line; linear to lanceolate, entire; usually exstipulate, and if stipulate then the stipules are
scarious (Paronychia); leaves highly reduced in arctic forms (Lychnis).
Inflorescence Flowers either solitary terminal (Arenaria) or arranged in dichasial cymes (cincinus-
type).
Flower Bracteate, bracteolate (Spergula, Fig. 21.21A), complete, hermaphrodite but rarely unisexual
(Lychnis); actinomorphic; hypogynous but perigynous in Arenaria; pentamerous but tetramerous in
Sagina; many bracts appear like epicalyx in Dianthus.
Calyx 5 sepals, polysepalous (Fig. 21.21A) or united into a tube at the base (Fig. 21.21B); rarely 4
sepals (Sagina); quincuncial or imbricate and persistent.
Corolla 5 petals, free; sometimes 4, rarely minute or none (Sagina); often notched (Cerastium) at the
tip; differentiated into a claw and limb; caryophyllaceous; petals deeply bifid in Stellaria media (Fig.
21.23); a ligule-like outgrowth is present on the petals in some genera of subfamily Silenoideae.
Androecium Stamens typically 5 or 10, or as many as petals, or twice as many as petals; arranged
in 1 or 2 whorls; obdiplostemonous i.e. stamens of outer whorl lie opposite the petals while that of
inner whorl alternate with the petals; polyandrous or basally connate; staminodia sometimes pres-
ent (Stellaria media, Arenaria); stamens of outer whorl are absent in Arenaria; in Stellaria media
some or all the stamens of outer whorl and even 2–3 of inner whorl are missing or present in the
form of staminodes; only 8 stamens in 2 whorls in Sagina; 5 stamens of inner whorl are absent in
Paronychia; anthers 2-celled, longitudinally dehiscent; introrse.
Floral developmental studies, however, indicate that instead of real obdiplostemony, Caryophyllaceae
show false obdiplostemony created by the mechanical pushing-out of the inner whorl of stamens.
A B
Fig. 21.21 Caryophyllaceae: Floral diagrams—A: Spergula arvensis and B: Silene conoidea.
Gynoecium Carpels 2 (Dianthus, Saponaria), 3 (Silene) or 5 (Lychnis); syncarpous; superior but

slightly inferior in Arenaria; unilocular; numerous ovules; free-central placentation; single basal
ovule in Paronychia; styles and stigmas as many as the carpels.
Fruits and Seeds Fruit is usually a dry capsule opening by valves or teeth; rarely in achene or utricle;
very rarely a berry (Cucubalus). Seeds with usually curved embryo around the perisperm.
Pollination and Dispersal Pollination is entomophilous. Seeds can not escape from fruit until shaken
by wind or animals.
General Floral Formula Br, Brl, ≈, , K5, C5, A5– 10, G(2 – 5).

• Ornamental plants Amongst several ornamentals of the family, some important ones include
Dianthus caryophyllus (carnation), D. berbatus (sweet William), D. plumarius (pink), D.
sinensis (rainbow pink), Gypsophila paniculata (baby’s breath), Silene pendula (catchfly),
Lychnis alba (white campion), Saponaria vaccaria (cowcockle), Arenaria grandiflora,
Cerastium arvense and species of Petrocoptis.
• Medicinal value (i) Dianthus anatolicus is used to cure intermittent fever; (ii) D. sinensis
is used as a diuretic and also against gonorrhoea; (iii) Lychnis coronaria is used against
diseases of liver and lungs; (iv) Silene apetala is useful in eye troubles; (v) Spergula arven-
sis seeds are used in lung tuberculosis; (vi) Saponaria vaccaria sap is used against urinary
bladder diseases; (vii) Stellaria media leaves are used against inflammations of digestive
and respiratory tracts; (viii) The extract of the entire plant of Stellaria semivestita has anti-
cancerous properties.
• Fodder for cattles is obtained from Spergula arvensis and several other species of this
family.
252 Plant Taxonomy
• Roots of Saponaria vaccaria (syn. Vaccaria pyramidata), Gypsophila, etc. contain saponin
and used for washing wood and silk.
• Perfume is obtained from the flowers of Dianthus caryophyllus.
• Drymaria cordata is used to check soil erosion.
• Troublesome weeds of the family include Stellaria media, Cerastium, Silene conoidea,
Saponaria vaccaria and Polycarpaea.
ovary wall
ouvles
locules
T.S. Ovary T.S. Ovary
(at the base) (higher up)
flower Floral Diagram
stamen
petal
pedicel
A Flower
stem stigma
stigma stamen
style
leaf
style
ovary
ovary
pedicel sepal
Flowering Branch petal pedicel
Gynoecium L.S. Flower
Fig.21.22 Dianthus caryophyllus L.

This family belonging to order Caryophyllales (Bentham and Hooker) is usually classified into
two subfamilies viz. Alsinoideae (with free sepals) and Silenoideae (with fused sepals). Tutin et al.
(1964) divided it in their Flora Europaea into three subfamilies, i.e. Alsinoideae, Paronychioideae
and Silenoideae. Cronquist (1981) placed it under order Caryophyllales along with 11 more families
whereas Thorne (1983) discussed Caryophyllaceae under order Chenopodiales of Centrospermae.
Usually, Caryophyllaceae is considered to have been derived from Ranales. However, Eichler
considered it to have originated from Phytolaccaceae. Dickson (1936) believed that Caryophyllaceae
probably originated from Geraniaceae.

1. Dianthus caryophyllus (Carnation, Fig. 21.22)
Habit: Annual, ornamental herb. Stem: Aerial, erect, solid, herbaceous. Leaf: Simple, exstipu-
late, opposite decussate, sessile, united at the base, lanceolate, entire, acute, unicostate reticulate.
Inflorescence: Solitary terminal or dichasial cyme. Flower: Multibracteate, bracts appear like epica-
lyx; pedicellate, complete, hermaphrodite, actinomorphic, hypogynous, pentamerous. Calyx: 5 sepals,
gamosepalous, tubular, quincuncial or valvate. Corolla: 5 petals, polypetalous, caryophyllaceous,
twisted. Androecium: 10 stamens, arranged in 2 whorls of each, polyandrous, obdiplostemonous;
anthers dithecous, dorsifixed, introrse. Gynoecium: Bicarpellary, syncarpous, superior; ovary uni-
locular at base but bilocular at top; ovules many, free-central placentation; styles 2, long; stigma
simple. Fruit: Capsule.
Floral Formula: Br, ≈, , K(5), C5, A5 + 5, G(2).
2. Stellaria media (Chickweed, Fig. 21.23)
Habit: Annual, herbaceous common weed. Leaf: Simple, opposite decussate, exstipulate, ovate or
oblong; entire, acute, unicostate reticulate. Inflorescence: Dichasial cyme. Flower: Bracteate, ebrac-
teolate, rarely 2 bracteoles are present; pedicellate, complete, actinomorphic, hermaphrodite, pentam-
erous, hypogynous. Calyx: 5 sepals, free, quincuncial, green. Corolla: 5 petals, free, petals deeply
bilobed appearing as if they are 10; imbricate or valvate. Androecium: Stamens 10 or reduced to
5 or 8, leaving the rest abortive in the form of staminodes; present in 2 whorls; obdiplostemonous;
anthers dithecous, basifixed, introrse. Gynoecium: Tricarpellary, syncarpous, superior, unilocular,
numerous ovules, free-central placentation; styles 3 but reduced; stigmas 3. Fruit: Capsule.
Floral Formula: Br, ≈, , K5, C5, A10 or 5 – 8, G(3).
21.16 PORTULACACEAE PURSLANE FAMILY

Polypetalae, Thalamiflorae, Caryophyllineae.

Annual or perennial herbs with usually fleshy leaves; scarious stipules; sepals 2, persistent; petals
4–6; ovary unilocular; basal placentation.

Portulaca, Talinum.
254 Plant Taxonomy
stigma stigma
stamen
ovary wall
sepal locule
style ovule
petal
ovule
ovary
ovary
thalamus
pedicel T.S. Ovary
L.S. Flower Gynoecium
flower
stem
leaf
Fig. 21.23 Stellaria media L.

Portulacaceae is represented by about 20 genera and 500 species (Jones and Luchsinger, 1987).
Though cosmopolitan, the members are mainly distributed in western North America and southern
South America. Only 7 species of Portulacaceae have so far been reported from India. Some larger
genera of the family along with their number of species in bracket include Portulaca (purslane,
200), Calandrinia (red maids, 150), Anacampseros (70), Talinum (50), Claytonia (spring beauty,
35), Lewisia (20) and Montia (15).

General Habit Mostly annual, sometimes perennial herbs or suffrutescent shrubs.
Root Branched, tap root; fleshy in Lewisia.
Leaves Alternate or opposite; often succulent, simple; stipulate, stipules sometimes in the form of
hairs or scales; exstipulate in Claytonia.
Inflorescence Usually cymose, or racemose, or solitary flowers.
Flower Bracteate or ebracteate, bisexual, actinomorphic, showy, complete, hypogynous or
perigynous.
Calyx Sepals usually 2, often persistent, free or united at the base. Some regard sepals as bracteoles,
and according to them the flowers have petaloid perianth.
Corolla 4 to 6 petals, free or united at the base; caducous; usually imbricate, but valvate in
Claytonia; showy.
Androecium 4 to many, free stamens; usually 5 stamens, opposite to petals; anthers dithecous,
introrse, dehiscing longitudinally; stamens 2 in Montia and 8–10 in Portulaca oleracea.
Gynoecium Usually tricarpellary, syncarpous, superior but partly inferior in Portulaca; unilocular;
2 to many campylotropous ovules on a central basal placenta; styles 2–5; stigmas 2–5 or as many
as carpels.
Fruit and Seed Fruit usually a dehiscent capsule; rarely a nut and indehiscent. Seeds with embryo
curved around the perisperm.
Pollination and Dispersal Pollination entomophilous and seeds are dispersed by animals or rains.
General Floral Formula Br or Ebr, ≈, , K2 or (2), C4– 6, A4– μ, G(3).

Portulacaceae is of little economic importance except of some well-known garden ornamentals,
such as Portulaca grandiflora for its pink, red, yellow or white flowers, P. afra for its rose-coloured
flowers, and P. oleracea for its showy flowers. Other ornamental plants of the family include
several species of Calandrinia, Talinum and Lewisia. Portulaca oleracea is sometimes used as
a pot herb and in salads and also in the ailments of kidney and urinary bladder. P. quadrifida
proves effective in cough and asthma.

Portulacaceae belongs to order Caryophyllineae (Bentham and Hooker). Cronquist (1981) discussed
it under Caryophyllales. However, Thorne (1983) placed it under suborder Portulacineae of order
Chenopodiales of Centrospermae.
Portulacaceae is related closely to the Cactaceae and Aizoaceae of order Caryophyllales (Hickey
and King, 1988).
256 Plant Taxonomy
Recent anatomical findings of the family suggest that its 2 sepals are actually the bracts and its
petals are thus the sepals.

1. Portulaca oleracea L. (Common Purslane; Fig. 21.24)
Habit: Herb with prostrate or ascending branches. Stem: Aerial, branched, succulent. Leaves: Simple,
opposite or alternate, stipulate, sessile or subsessile, apex obtuse or truncate, entire. Inflorescence:
Flowers solitary or in terminal clusters. Flower: Ebracteate, complete, hermaphrodite, actinomorphic,
perigynous. Calyx: 2 sepals, free, anterio-posterior. Corolla: 5 petals, free, quincuncial. Androecium:
8–12 stamens, usually 10, polyandrous, in one whorl, adnate, dithecous, introrse. Gynoecium: Tri- to
pentacarpellary, syncarpous, half-inferior, unilocular, many ovules, free-central placentation; styles
3–5. Fruit: Capsule.
Floral Formula: Ebr, ≈, , K2, C5, A8 – 12 or 10, G (3– 5)–.
petal
stigma
style
stamen
A Stamen
sepal
ovary
L.S. Flower
stem
roots
Flowering Plant
Fruit T.S. Ovary
Floral Diagram
Fig. 21.24 Portulaca oleracea L.

21.17 MALVALES
1. Flowers bisexual and actinomorphic, and only rarely zygomorphic.
2. Flowers usually pentamerous with their calyx valvate.
3. Petals usually as many as sepals; epicalyx usually present.
4. Stamens numerous and usually monadelphous.
5. Ovary tri- to multicarpellary.
6. Placentation usually axile.
Cohort Malvales of Bentham and Hooker includes 3 orders = (families) namely Malvaceae,
Sterculiaceae and Tiliaceae. Engler and Diels, however, treated the order Malvales to be composed
of 4 suborders and 8 families as under:
1. Elaeocarpineae: Elaeocarpaceae
2. Chlaeneae: Chlaenaceae
3. Malvineae: Tiliaceae, Malvaceae, Bombacaceae, Sterculiaceae
4. Scytopetalineae: Scytopetalaceae
Hutchinson, however, included only Malvaceae under order Malvales, and treated Sterculiaceae,
Bombacaceae and Tiliaceae in a new order Tiliales.
Tiliaceae, Malvaceae, Sterculiacene and Bombacaceae have been treated in this text.
21.18 MALVACEAE MALLOW FAMILY

Polypetalae, Thalamiflorae, Malvales.

Shrubs or herbs, frequently with stellate trichomes; sepals persistent; monadelphous stamens; anthers
monothecous; carpels 5 or more.

Gossypium, Hibiscus, Althaea, Malvastrum, Sida, Malva, Abutilon.

A family of about 85 genera and 1,000 to 1,500 species, Malvaceae are distributed widely in tropi-
cal and temperate regions. 22 genera and about 125 species of Malvaceae have so far been reported
from India. Some of the larger genera, along with their common names and/or number of reported
species in parenthesis, include Hibiscus (rose mallow, 300), Sida (200), Pavonia (200), Abutilon
(Indian mallow, 100), Alcea (60), Malva (mallow, 40), Lavatera (25), Gossypium (cotton, 20), and
Althaea (12).
258 Plant Taxonomy
The family is recognized by Hibiscus rosa-sinensis (rose of China) because of its beautiful large
flowers and hundreds of its known cultivated varieties. Cotton (Gossypium), the most important plant
of this family from the commercial viewpoint, has been cultivated in India since last 5,000 years.

General Habit Mostly annual (Althaea, Malva) or perennial herbs, sometimes growing very tall
(Lavatera); shrubs (Gossypium, Hibiscus rosa-sinensis); or trees (Hibiscus elatus, Thespesia popul-
nea, Kydia); often with mucilaginous sap.
Root Profusely branched, tap roots.
Stem Erect, branched, sturdy; young portions often covered with stellate trichomes; decumbent in
Malva rotundifolia.
Leaf Simple, often palmately lobed and palmately veined; alternate; stipulate, stipules often decidu-
ous; petiolate; ovate, cordate or reniform; margin toothed or entire; venation reticulate and multi-
costate divergent.
Inflorescence Usually cymose or flowers solitary in the leaf axil (Hibiscus rosa-sinensis); raceme
in Althaea rosea.
Flower Bracteate; bracteolate, bracteoles often form calyx-like epicalyx; complete, bisexual, actino-
morphic, hypogynous, pentamerous; flowers rarely unisexual and the species are dioecious (Napaea);
variously coloured.
Epicalyx Outside the calyx, several bracteoles often form a calyx-like whorl called epicalyx
(Fig. 21.27). Epicalyx consists of 3 bracteoles (Malvastrum tricuspidatum, Malva), or 5–9 brac-
teoles (Althaea rosea), or several bracteoles (Hibiscus radiatus); bracteoles are free (Malvastrum)
or fused (Althaea); epicalyx or involucre of bracteoles is absent in Abutilon (Fig, 21.26) and Sida
(Fig, 21.25).
Calyx 3 to 5 but usually 5 sepals, free or basally connate,
valvate; tubular in Hibiscus.
Corolla 5 petals, free but basally adnate to the staminal col-
umn; usually twisted but sometimes imbricate; showy and vari-
ously coloured.
Androecium Numerous stamens, monadelphous; anthers 1-celled
or monothecous; reniform; extrorse; dehiscence longitudinal;
filaments of anthers fuse to form a staminal column around
the ovary.
Gynoecium A compound pistil of 1 to many carpels (often
pentacarpellary, syncarpous); ovary superior; 2 to 5 or more
locules, rarely 1 locule; 1 to numerous ovules in each locule;
Fig. 21.25 Floral diagram of axile placentation; style 1 and branched above, or as many as
Sida cordifolia. the carpels; stigmas as many or twice as many as carpels, dis-
coid or capitate.
stigmas
petal
sepal
stamen
ovules
ovary
thalamus
pedicel
L.S. Flower
Floral Diagram
anther
lobe
connective
filament
fruit
Stamen flower
Fruit
leaf
stigmas
stem
style ovary
sepal
pedicel
Gynoecium Flowering Branch
Fig. 21.26 Abutilon indicum Sweet.
Fruit and Seed Fruit usually a capsule (Gossypium) or schizocarp (Malva), or rarely a berry
(Malvaviscus) or samara. Seeds with curved embryo and scanty or no endosperm.
Pollination and Dispersal Majority of Malvaceae are insect pollinated mainly because of their large-
sized flowers, brightly coloured corolla and extrafloral nectaries (in some). Dispersal of seeds takes
place mainly by wind (Gossypium), or sometimes by water (Malva), or by animals because of the
presence of hooked spines (Urena lobata).
General Floral Formula Br, ≈, , Epik3 – 9 or (3– 9), K(5), C5 or (5), A (μ), G(2 – 5) or (1 – μ).
260 Plant Taxonomy

• Cotton: Cotton, manufactured from the dense mass of long hairs attached to the seeds of
several species of Gossypium, is the most important product of this family. Cotton comes
from Gossypium barbadense, G. hirsutum, G. arboreum and G. herbaceum.
• Other Fibres: Hemps, which are the bast fibres (i.e. obtained from the phloem fibres of
the stem), are obtained from Hibiscus elatus (Cuba bast), H. sabdariffa (Rozella hemp), H.
cannabinus (Deccan hemp), Sida rhombifolia (Queensland hemp), etc. Bags, ropes, cords,
etc. are made from the fibres obtained from Abutilon asiaticum, A. theophrasti, Hibiscus
falcatus, H. tiliaceous, Abelmoschus moschatus, Urena lobata, etc.
• Oils: (i) Cotton seed oil, which is edible and also used for the manufacture of soaps,
lubricants, etc., is obtained from the seeds of Gossypium. The oil cake is an excellent cattle
feed. (ii) An essential oil, used in perfumery, is obtained from the seeds of Abelmoschus
moschatus. (iii) A fatty oil, obtained from the seeds of ‘Patsan’ (Hibiscus cannabinus), is
used in the manufacture of paints, varnishes and linoleum, and its refined oil is edible.
• Ornamentals: Malvaceae are known universally for their ornamental plants, chief among
which include Hibiscus rosa-sinensis (China rose or shoe flower), H. mutabilis (cotton rose),
H. Schizopetalus, H. syriacus (Rose of Sharon), H. sabdariffa, H. collinus, Althaea rosea
(Hollyhock), Malva sylvestris (Mallow), Malvaviscus arboreus, Thespesia populnea, etc.
• Vegetables: (i) Lady’s finger or okra, a famous vegetable, is the fruit of Abelmoschus
esculentus (vern. Bhindi); (ii) a delicious ‘chutney’ is prepared from the sepals of Hibiscus
cannabinus and H. sabdariffa; (iii) flower buds of Hibiscus syriacus are acidic and used
as vegetable.
• Timber: Wood of Thespesia populnea is used in making boats while that of Hibiscus elatus
is utilized in furniture making.
• Medicinal Value: (i) Roots of Abutilon indicum are used against fever while that of
Abelmoschus moschatus in stomach ache, and that of Althaea officinalis and Hibiscus rosa-
sinensis against cough; (ii) roots of Althaea rosea are used in the treatment of dysentry; (iii)
bark of Gossypium is used for stopping haemorrage in ladies, specially after child birth; (iv)
roots of Malva verticillata are used against whooping cough; (v) leaves and seeds of Malva
sylvestris are used in fever; (vi) roots of Malachra capitata are used in rheumatism; (vii)
roots and bark of Urena repanda are used to cure hydrophobia, a dreaded disease caused
by bite of mad dog.
• Some Other Uses: (i) ‘Hina’, a well-known Indian perfume, is obtained from the roots
of Pavonia odorata; (ii) toys, pencils and matchsticks are manufactured from the wood of
Thespesia populnea; (iii) a blue dye is obtained from the leaves of Althaea rosea; (iv) a
well-known drink is prepared from the calyx of Hibiscus sabdariffa.

Malvaceae, placed under order Malvales by Bentham and Hooker, has been retained under the same
order (Malvales) by almost all recent taxonomists including Cronquist (1981) and Thorne (1983).
Schumann (1964) divided it into 4 tribes viz. Malopeae, Malveae, Ureneae and Hibisceae.
stigma
stigmas
style
anthers
ovary staminal
tube
thalamus
Gynoecium
Essential Organs
stigma
Floral Diagram
stamens
staminal
tube
stigma
corolla stamen
calyx
staminal
tube
ovary ovule
calyx
epicalyx
pedicel
L.S. Flower
anther
lobe
leaf
filament
stem
Stamen
Flowering Branch
Fig. 21.27 Hibiscus rosa-sinensis L.

262 Plant Taxonomy
Malvaceae are related to Bombacaceae but members of the former bear monadelphous androecium
while that of the latter bear polyadelphous condition.
In several respects Malvaceae also show affinities with Sterculiaceae and Tiliaceae. However,
the former bears monothecous anthers whereas the members of Sterculiaceae and Tiliaceae possess
dithecous anthers.

1. Abutilon indicum Sweet. (Vern. Kanghi, Fig. 21.26)
Habit: Annual herb. Stem: Erect, herbaceous, branched, solid, hairy. Leaf: Simple, alternate, petiolate,
stipulate, serrate, acute, multicostate reticulate. Inflorescence: Solitary axillary. Flower: Bracteate,
ebracteolate, pedicellate, complete, hermaphrodite, actinomorphic, pentamerous, hypogynous. Calyx:
5 sepals, gamosepalous, valvate. Corolla: 5 petals, free but slightly united at the base and adnate to
staminal tube, twisted. Androecium. Numerous monadelphous stamens forming staminal tube around
style, monothecous, extrorse. Gynoecium: Polycarpellary, syncarpous, superior, multilocular, one
ovule in each locule, axile placentation; style long; stigmas as many as carpels. Fruit: Capsule.
Floral Formula: Br, Ebrl, ≈, , K(5), C5 or (5), A (μ), G(μ).
2. Hibiscus rosa-sinensis L. (Chinese-rose or Gurhal, Fig. 21.27)
Habit: Typical ornamental shrub of the family. Leaf: Simple, alternate, petiolate, stipulate, ovate,
serrate, acute. Inflorescence: Solitary axillary. Flower: Ebracteate, complete, hermaphrodite, acti-
nomorphic, large, red, showy. Epicalyx: 5–8 bracteoles around the calyx form epicalyx. Calyx: 5
sepals, fused, valvate. Corolla: 5 petals, free but adnate to the staminal tube, twisted, red, petals
with sinuous margins. Androecium: Same as that of Abutilon. Gynoecium: 5 fused carpels, superior
ovary, pentalocular, one to many ovules in each locule, axile placentation; styles 5, long, united below
and passing through the staminal tube; stigmas 5 and discoid. Fruit: Capsule.
Floral Formula: Ebr, ≈, , Epik5 –8, K(5), C5, A(μ), G(5).
21.19 STERCULIACEAE STERCULIA FAMILY


Generally shrubs or trees with stellate hairs; leaves alternate, stipulate; sepals and petals 5, some-
times petals modified or absent; stamens generally in 2 whorls, of which outer whorl is reduced to
staminodes; gynoecium generally pentacarpellary, syncarpous; some bear androgynophore.

Abroma, Helicteres, Sterculia, Theobroma, Pterospermum, Melochia.
21.19.4 Size and Distribution

Represented by about 60 genera and 700 species, Sterculiaceae are distributed exclusively in tropi-
cal and subtropical parts of the world. About 18 genera and 90 species of this family have been
reported from India.

General Habit Usually shrubs or trees with soft wood; rarely herbs with mucilage (Waltheria,
Melochia); sometimes lianous (Buettneria).
Stem Well-branched, woody, with its younger parts covered with stellate hairs.
Leaf Simple, alternate or subopposite; stipulate, stipules caducous; pulvinous petiole; entire or pal-
mately lobed (Fig. 21.28) or compound; covered with stellate hairs.
Inflorescence Axillary or terminal compound cymose (Fig. 21.28); sometimes cauliflorous
(Theobroma) i.e. flowers borne on stem.
Flower Bracteate, bracteolate (Melochia), bisexual, sometimes unisexual (Cola); actinomorphic,
hypogynous, pentamerous; androgynophore present in some (Mansonia).
Calyx 3–5 sepals, free or basally connate, hairy, petaloid (Sterculia), valvate.
Corolla 5 petals, but occasionally absent in Sterculia (Fig. 21.28), polypetalous; hooded in
Theobroma; twisted or imbricate.
Androecium Usually 5 to 10 stamens, free or monadelphous; in two whorls, of which outer whorl of
stamens is antisepalous and reduced to staminodes or scales or even absent, and filaments of inner
whorl of stamens connate (monadelphous); anthers 2-celled; dehiscence longitudinal; stamens are 20
in Pentapetes and Pterospermum, of which 5 antipetalous stamens are reduced to staminodes.
Gynoecium Bi- to pentacarpellary, syncarpous, superior ovary raised over an androgynophore in
Cola, Mansonia; bi- to pentalocular; 2 to many ovules in each locule; axile placentation; mono-
carpellary and unilocular condition is seen in Waltheria; style simple or divided into lobes, each
lobe terminates into a stigma.
Fruit and Seed Fruit is a capsule (Pterospermum), follicle (Helicteres), or drupe (Theobroma). Seeds
are arillate, often winged, with straight or curved embryo and abundant endosperm.
Pollination and Dispersal Pollination generally entomophillous, and seeds are dispersed generally by
wind and also sometimes by animals.
General Floral Formula Br, Brl, ≈, , K3–5 or (3–5), C5, A5+(5), G (2–5).

• Chocolate, cocoa and cocoa-butter are made from the seeds of Theobroma cacao, which is
rich in alkaloids thein and theobromine.
264 Plant Taxonomy
Flower Flower
seeds (side view) (top view)
Dehiscing
Fruit
flowers
Floral Diagram
A Leaf
Flowering Branch
Fig. 21.28 Sterculia villosa Roxb.
• Soft drinks are prepared from the cola nuts (Cola acuminata), which contain thein and
caffeine.
• Fruits of Guazuma and seeds of Sterculia are edible whereas valuable fibres are obtained
from the bark of Sterculia urens, S. villosa, Helicteres isora and Guazuma ulmifolia.
• Wood of Heritiera minor is utilised for domestic purposes.
• Flowers of Pterospermum acerifolium are used as disinfectant and insect-repellant.

• Ornamental plants of Sterculiaceae, grown usually in the gardens, include species of Abroma,
Brachychiton, Dombeya, Firmiana, Guazuma, Pentapetes, Pterospermum and Reevesia.
• Useful timber, utilized for toys, match-sticks, tea boxes, etc. is obtained from species of
Eriolaena candllei, Heritiera littoralis, Petrospermum diversifolium, Sterculia foetida and
S. villosa.
• Gum of commercial importance is obtained from several species of Sterculia.
• Diabetes is cured by the juice of the roots of Helicteres isora, whereas roasted seeds of
Guazuma ulmifolia are used in stomach troubles. Fruits and seeds of Sterculia balanghas
have laxative properties.

Placed under order Malvales by most of the taxonomists (Bentham and Hooker, Engler and Prantl,
Takhtajan, Cronquist, Thorne, etc.), Sterculiaceae are treated under Tiliales by Hutchinson. This
family differs from Malvaceae and Tiliaceae in the absence of epicalyx and presence of definite
number of stamens. Because of the presence of characters, such as stellate hairs, tendency of uni-
sexuality, and arillate seeds, Sterculiaceae show affinities with Euphorbiaceae. However, Sterculiaceae
are generally considered to have been derived from the same stock as the Tiliaceae.
21.20 TILIACEAE LIME FAMILY OR BASSWOOD FAMILY


Woody trees or shrubs with strong phloem fibres, rarely herbs; leaves alternate, simple, stipulate;
flowers bisexual, actinomorphic, usually pentamerous, arranged in cymes or panicles; stamens 5 to
many, often polyadelphous; axile placentation; fruit drupe or capsule.

Tilia, Corchorus, Triumfetta, Grewia.

Tiliaceae includes about 50 genera and 550 species distributed in tropical and temperate regions,
mainly in South East Asia and Brazil. Tilia is mainly distributed in temperate regions. About 15
genera and over 100 species of Tiliaceae are found in India. Some of the larger genera of the fam-
ily, along with the number of their approximately reported species in parenthesis, are Grewia (150),
Triumfetta (150), Corchorus (100), Tilia (50) and Sparmannia. Grewia subinaequalis (Phalsa),
Corchorus capsularis (Jute) and Elaeocarpus ganitrus (Rudraksh) are some of the well-known spe-
cies of Tiliaceae.
266 Plant Taxonomy

General Habit Usually trees (Tilia) or shrubs (Grewia), rarely herbs (species of Triumfetta,
Corchorus) or climbers (Plagiopteron).
Stem Woody, branched, contains strong phloem fibres; often with stellate pubescence.
Leaf Usually alternate, simple, stipulate; rarely opposite (Plagiopteron); petiolate; margin dentate
or serrate; covered with stellate or branched hairs; leaf blade often oblique with larger side towards
the branch.
Inflorescence Cymose; rarely axillary racemes (Elaeocarpus); greatly condensed dichasial cymes
in Corchorus.
Flower Bracteate; bracteolate (Fig. 21.29), bracte-
oles often appear like epicalyx; pedicellate, complete,
hermaphrodite but rarely unisexual; actinomorphic,
hypogynous; usually pentamerous; sometimes with
androgynophore (Duboscia, Grewia).
Calyx Usually 5 sepals, rarely 3 or 4; polysepal-
ous but gamosepalous in Chartacalyx; generally
hairy, valvate; nectar-secreting glands at the base
of sepals in Tilia.
Corolla Usually 5 petals, rarely less or even absent
(Chartacalyx); polypetalous, imbricate; usually yel-
low coloured but rarely sepaloid (Elaeocarpus);
often glandular at base.
Androecium Usually 10 or more stamens, free or
often united by the basal part of their filaments
into 5 or 10 bunches (polyadelphous); some of
the stamens may modify into petaloid staminodes
Fig. 21.29 Floral diagram of Tilia. (Pentace); anthers 2-celled, dehiscing by longitudi-
nal slits or apical pores; in Grewia androgynophore
is present.
Gynoecium Bi- to multicarpellary, syncarpous, superior; 2 to many loculed, one to many ovules
in each locule, axile placentation; style one dividing into as many stigmas as number of carpels; in
Elaeocarpus, septa in the ovary get dissolved to make it unilocular.
Fruit and Seed Fruit variable, fleshy or dry, and dehiscent or indehiscent; fruit usually a capsule
(Corchorus, Fig. 21.30) or schizocarp. Seeds with straight embryo; endosperm copious or scanty.
Pollination and Dispersal Pollination entomophilous, usually by flies or bees; entomophily is favoured
because of protandry and presence of nectar or scent secreted in the spoon-like glands at the base
of the sepals in some (Tilia).
General Floral Formula Br, Brl, ≈, , K5, C5, A10 – μ, G(2 – μ).
stigma
petal
stamens
style
ovary
sepal
ovules
pedicel
L.S. Flower
flower
bud
Floral Diagram
fruit locule
ovule
stipule
ovary
leaf wall
anther lobes
T.S. Ovary
connective
stem
filament
Stamen Flowering Branch
Fig. 21.30 Corchorus aestuans L.

• Jute Two species of Corchorus (C. capsularis and C. olitorius) are grown widely in warm
areas as the source of the jute fibre. Jute is obtained from the bast or phloem fibres. It is
cultivated extensively in Bangladesh, and West Bengal in India.
• Timber Several Tilia species are grown for their timber in temperate regions. Species of
Berrya, Grewia, Elaeocarpus and Erinocarpus are also grown for similar purposes.
• Ornamentals Several Tiliaceae are planted for ornament in parks, large gardens and
on roadsides because of their fragrant flowers. Common among them are Sparmannia
africana (House Lime), Tilia americana (Basswood), T. europaea (European Linden),
268 Plant Taxonomy
T. tomentosa (White Linden), Grewia elastica, G. serrulata, Elaeocarpus ganitrus and

Muntingia calabra.
• Edibles Fruits of Grewia asiatica syn. G. subinaequalis (Phalsa), G. tenax, G. tilifolia and
G. villosa are edible.
• Medicinal Value (i) Dried leaves and decoction of roots of Corchorus capsularis and
C. olitorius are used to cure dysentery and diarrhoea; (ii) root bark of Grewia asiatica
proves effective in rheumatism while its fruits help in digestion; (iii) G. microcos plants
are used in eczema; (iv) roots of G. sclerophylla are prescribed in cough; (v) wood of G.
tiliaefolia is emetic and taken as an antidote to opium poisoning; (vi) roots of G. villosa are
used in diarrhoea; (vii) leaves, flowers and fruits of Triumfetta bartramia are used against
gonorrhoea.
• Fodder Leaves of Grewia glabra, G. oppositifolia, G. sapida and G. tenax are used as
fodder for cattles.

Tiliaceae belongs to order Malvales according to most of the botanists (Bentham and Hooker, 1862–
1883; Rendle, 1925; Cronquist, 1981; and Thorne, 1983). However, Hutchinson (1959) placed this
family under order Tiliales. Tiliaceae is closely allied to Malvaceae. However, members of Tiliaceae
possess dithecous anthers and their stamens are nearly distinct whereas those of Malvaceae have
monothecous anthers and their androecium is monadelphous. Tiliaceae is similar to Sterculiaceae
in habit, leaves and stipules but differs in possessing indefinite androecium. According to some
botanists, Tiliaceae is allied to some genera of Euphorbiaceae which have free stamens attached
upon a gynophore.

1. Corchorus aestuans Linn. syn. C. acutangulus Lamk. (Fig. 21.30)
Stem: Branched, aerial, herbaceous, solid, hairy, dark red. Leaf: Simple, alternate, petiolate, stipu-
late, ovate, serrate, acute, unicostate reticulate. Inflorescence: Axillary cyme. Flower: Bracteate,
bracteolate, pedicellate, complete, hermaphrodite, actinomorphic, hypogynous, pentamerous. Calyx:
5 sepals, free, valvate. Corolla: 5 petals, polypetalous, valvate, yellow. Androecium: 10–20 stamens
in 2 to 4 whorls of 5 stamens in each whorl, polyandrous, dithecous, basifixed or dorsifixed, introrse.
Gynoecium: Tricarpellary, syncarpous, superior ovary, trilocular, 2 ovules in each locule, axile pla-
centation; style single; stigma trifid, each stigmatic lobe is again bifurcated. Fruit: Capsule.
Floral Formula: Br, Brl, ≈, , K5, C5, A10–20, G(3).
21.21 BOMBACACEAE BOMBAX FAMILY

Treated as a subfamily of Malvaceae by Bentham and Hooker.

Exclusively arborescent or large-sized trees; simple or palmately compound leaves; stipules caducous;
large-sized flowers; anthers dithecous or monothecous; pollen grains always smooth; fruit capsule.

Adansonia, Bombax, Ceiba, Salmalia, Cullenia.

A small family of about 22 genera and 180 species, Bombacaceae are mainly distributed in tropical
regions. Bombax malabaricum syn. Salmalia malbarica (Simbal) is very common in India whereas
Durio zibethinus is indigenous to Malayan archipelago.

General Habit Tall trees with large, thick and spreading branches (Adansonia, Ceiba pentandra).
Stem Aerial, erect, branched, tall, thick; a few have extremely light wood (Ochroma lagopus).
Leaf Alternate, simple (Cullenia) or palmately compound; stipulate, stipules deciduous; petiolate.
Inflorescence Solitary axillary or in axillary clusters.
Flower Bracteate, rarely ebracteate, pedicellate, complete, bisexual, actinomorphic, large, showy,
hypogynous.
Calyx 5 sepals, gamosepalous, valvate (Fig. 21.31); leathery in texture.
Corolla 5 petals, polypetalous, imbricate or twisted; petals rarely absent (Cullenia excelsa).
Androecium Stamens 5 (Ceiba pentandra) to numerous (Salmalia malabarica), free or monadel-
phous (Adansonia, Ceiba) or polyadelphous (Bombax malbaricum; Fig. 21.31); staminodes often
present; anthers 1-, 2- or more-celled containing smooth pollen grains; stamens very long in Pachira
insignis; dehiscence longitudinal; extrorse.
Gynoecium 2–5 carpels, syncarpous, ovary superior, 2- to 5-locular, 2 or more ovules in each locule,
axile placentation; style long and simple; stigmas 1 to 5.
Fruit and Seed Fruit capsule. Seeds endospermic (Salmalia) or non-endospermic (Cullenia); bear
silky hairs in Ceiba pentandra.
General Floral Formula Br, ≈, , K(5), C5, A5 – μ or (5 – μ), G(2 – 5).

1. Fruits of Adansonia digitata provide us a good beverage.
2. Wooly outgrowth present on the pericarp of the fruits of Bombax ceiba is used for stuffing
pillows while the floss obtained from the seed hairs of Ceiba pentandra is the source of
‘Kapok’.
270 Plant Taxonomy
stigma
stamens
petal
style
sepal
Flower ovary
L.S. Flower
flower stigma
style
leaf
ovary
stem
Flowering Branch Gynoecium
Floral Diagram
Fig. 21.31 Bombax malbaricum D.C.
3. The flower buds and fruits of Bombax ceiba and Durio zibethinus are edible. Root decoc-
tion of D. zibethinus is used for fever.
4. World’s lightest commercial wood with a specific gravity of only 0.12 is obtained from
Ochroma lagopus and O. pyramidala.
5. Wood obtained from Cullenia excelsa is used for making pencils, plywood, packing cases,
etc.

In old classical literature of taxonomy, Bombacaceae has been considered as a taxon under family
Malvaceae. Bentham and Hooker also treated it as a subfamily of Malvaceae. However, in majority
of the recent taxonomic works including that of Cronquist (1981) and Thorne (1983), Bombacaceae
has been treated as an independent family of order Malvales similar to that of Malvaceae. Moreover,
Bombacaceae differs from that of Malvaceae in (i) being exclusively arborescent, (ii) often possess-
ing a prickly trunk, (iii) bearing the dithecous anthers in some and monothecous in others, and (iv)
always having smooth pollen grains.
In several anatomical and floral characters Bombacaceae shows close affinities with Malvaceae.
However, several of its genera show close relationship with Dilleniaceae on the basis of their stamen
morphology. Takhtajan (1969) and Cronquist (1981) show close relationship of Bombacaceae with
Malvaceae, Sterculiaceae and Tiliaceae.
21.22 GERANIALES
1. Stamens usually twice as many as sepals.
2. Disc present; usually disc is annular, adnate to stamens or reduced to glands.
3. Ovary multicarpellary and syncarpous.
4. Style often persistant in fruit.
5. Seeds usually non-endospermic or endosperm scanty.
According to Bentham and Hooker, Cohort Geraniales includes 11 orders (=families), of which
three major ones are Geraniaceae, Rutaceae and Meliaceae.
Engler and Diels divided order Geraniales into 6 suborders (Geraniineae, Malpighiineae,
Polygalineae, Dichapetalineae, Triococceae and Callitrichineae) and 21 families. Lawrence opined
that the “evidence from all fields of Botany indicates that the Geraniales (sensu Engler) are not a
natural taxon”.
Geraniaceae, Oxalidaceae, Rutaceae and Meliaceae are treated in this text.
21.23 GERANIACEAE GERANIUM FAMILY

Polypetalae, Disciflorae, Geraniales.

Herbs, sometimes suffrutescent; leaves palmately lobed or dissected; 5-merous flowers; stamens with
filaments united at base; beaked or lobed fruits with elastic dehiscent schizocarp.

Geranium and Pelargonium.

Geraniaceae is a family of about 11 genera (Jones and Luchsinger, 1987) and 780 species (Hickey
and King, 1988), which are cosmopolitan in distribution and found widely in temperate and tropical
regions. Only 3 genera and about 25 species have been reported from India, mainly from Himalayas.
Some of the larger genera along with their approximately reported number of species in parenthe-
sis include Geranium (400), Pelargonium (250), Erodium (90) and Monsonia (25). Well-known
‘geranium-oil’ is distilled from Pelargonium odoratissimum.

General Habit Mostly herbs, sometimes undershrubs; suffrutescent or arborescent; stems often fleshy
(Pelargonium); hairy.
272 Plant Taxonomy
stigma
style
ovary
Stamen of Outer Whorl

Flower disc
Gynoecium
receptive
stigma
style
mericarp
ovules
L.S. Gynoecium
Schizocarp Showing
Coiling Mericarps
rhizome
T.S. Ovary
Seed
Flowering Plant
Fig. 21.32 Geranium maculatum L.
Leaf Simple and lobed or dissected, or compound; opposite or alternate; stipulate; venation mostly
palmate.
Inflorescence Usually cymose, sometimes umbellate (Pelargonium), or flowers solitary.
Flower Bracteate, bracteolate, complete, hermaphrodite, actinomorphic but zygomorphic in
Pelargonium, hypogynous, pentamerous; often very attractive.
Calyx Usually 5 sepals, free or rarely united, imbricate; sepals 4 in Vivania and 8 in Dirachma;
persistent; in zygomorphic flowers the dorsal sepal is sometimes spurred and the spur is adnate to
the pedicel.
Corolla 5 petals, free, alternate with sepals, imbricate or twisted; rarely petals are 2, 4, 8 or 0;
often alternating with nectariferous glands.
Androecium 5 to 15 stamens; filaments often united at the base; in 1 to 3 whorls; obdiplostemonous;
staminodes often present (Erodium); anthers usually
versatile, 2-celled, longitudinally dehiscent; a nectary style
is present at the base of each stamen of inner whorl

rostrum
in Geranium. bristles
Gynoecium Pentacarpellary, syncarpous, superior
ovary, pentalocular, one to two ovules in each locule, empty ejected
mericarp seed
axile placentation; rarely 3 to 8 carpels; styles 3 to
5, slender and beak like; stigmas 3 to 5.
awn
Fruit and Seed Fruit capsular, dehiscing into 1 to
2-seeded 3 to 5 mericarps (Fig. 21.32), the styles central column
adhering to the ovarian beak. Seeds usually with
curved embryos; endosperm scanty or absent. remains of
stamens mericarp before
seed-ejection
Pollination and Dispersal Pollination mainly by
bees which visit the flowers in search of nectar. In sepal scar
Geranium, the dispersal of seeds takes place by the pedicel
awn, which suddenly curls upwards, throwing the
Fig. 21.33 Mechanism of seed dispersal
seeds at some distance from the plant (Fig. 21.33).
in Geranium pratense.
General Floral Formula Br, Brl, ≈, , K 5, C 5,
A(5–15), G(5).

• Ornamental value of Geraniaceae is because of the beautiful and attractive flowers of sev-
eral of its members, including Pelargonium hortorum, P. zonale, Geranium wallichianum,
G. pratense, Erodium cicutarium, etc.
• Well-known ‘geranium oil’, used in perfumery, is obtained from Pelargonium
odoratissimum.
• Roots and leaves of Geranium robertianum are used in the treatment of diarrhoea whereas
the rootstocks of G. wallichianum are used in eye troubles. Roots and leaves of G. nepalense
are used in the treatment of kidney troubles.
• Several species of Erodium and Geranium are weedy.
274 Plant Taxonomy
21.23.7 Systematics and Phylogeny

Geraniaceae belongs to order Geraniales, series Disciflorae and subclass Polypetalae of Dicotyledons
(Bentham and Hooker). Majority of taxonomists believe it to belong to order Geraniales. Thorne
(1983) placed Geraniaceae under suborder Geranineae of order Geraniales belonging to superorder
Geraniiflorae. Usually Geraniales are considered to have derived from Ranales. However, Takhtajan
(1969) stated that Geraniales are “clearly connected with Rutales, especially with the Rutaceae”.
21.24 OXALIDACEAE WOOD SORREL FAMILY

Treated as a tribe of Geraniaceae by Bentham and Hooker.

Usually palmately compound leaves with a sour taste due to the presence of oxalic acid; pentamer-
ous flowers with monadelphous stamens; styles 5; fruit loculicidal capsule.
21.24.3 Indian Genera

Oxalis and Biophytum.

The family includes only 8 genera and about 900 species, distributed mainly in tropical and sub-
tropical regions. Only a few species extend into temperate regions. Only 2 genera and about a dozen
species have so far been reported from India. Most common representative in India is Oxalis cor-
niculata (Indian Sorrel). Chief genera with their approximately reported species are Oxalis (800),
Biophytum (70), Averrhoa (2) and Eichleria (2). Biophytum sensitivum is sensitive to touch like that
of Mimosa pudica of Mimosoideae.

General Habit Mostly perennial herbs, rarely shrubs or trees (Averrhoa); often suffrutescent or
scapose and produce fleshy rhizomes or bulbous tubers; juice of the plants is sour because of the
presence of oxalic acid.
Leaf Alternate, pinnately or palmately compound; exstipulate; often long-petioled; leaflets obcordate
and characteristically folding at night; sometimes sensitive to touch (Biophytum sensitivum); leaves
sometimes replaced by phyllodes (Oxalis bupleurifolia).
Inflorescence Solitary or subumbellate, rarely racemose or cymose.
Flower Bracteate, bracteolate, complete, hermaphrodite, actinomorphic, pentamerous, hypogynous;
often on long peduncles.
Calyx 5 sepals, polysepalous (Fig. 21.34) or slightly united; imbricate or quincuncial, persistent.
Corolla 5 petals, free or slightly united at the base, short-clawed; imbricate or twisted.
Androecium Stamens 10, fused at the base (monadelphous), obdiplostemonous, introrse; rarely 5
stamens are without anthers.
sepals
style
stamen
stigma
carpels
Stamen ovule
(anterior view) stamen
Calyx
ovary
style
stigma
Style and Stigma

Flower with
Sepals & Petals
L.S. Ovary
Removed
petal
sepal
stamen
ovary
L.S. Flower T.S. Ovary
flower
Fruit
rhizome
roots
A Flowering & A Non-Flowering Plant
Floral Diagram
Fig. 21.34 Oxalis acetosella L.

276 Plant Taxonomy
Gynoecium Pentacarpellary, syncarpous, ovary superior, pentalocular, one or more anatropous ovules
in each locule, axile placentation; styles 5, free and persistent.
Fruit and Seed Fruit a loculicidal capsule, rarely a berry (Averrhoa). Seeds with straight embryo
and fleshy endosperm.
Pollination and Dispersal Pollination is usually by insects. Seeds are often discharged by the elastic
separation of fleshy aril of the seed coat from an inner harder layer.
General Floral Formula Br, Brl, ≈, , K5, C5, A(10), G(5).

The family is of little importance. The leaves of Oxalis (sour grass) are chewed by the children
for their pleasant sour taste due to oxalic acid. Tuberous roots of O. tuberosa are used as food in
India and South America. Leaves of O. acetosella (common wood sorrel) are used as salads and
stems of O. pescaprae as a vegetable. Ornamental plants of the family include Oxalis articulata
(bright pink or white flowers), O. deppei, O. hedysaroides, and both the species of genus Averrhoa
(evergreen trees). Biophytum sensitivum is sensitive to touch and grown in gardens as a curiosity,
as its leaves are sensitive like those of Mimosa pudica of Mimosoideae.

Bentham and Hooker treated Oxalidaceae only as a tribe Oxalideae under family Geraniaceae, belong-
ing to order Geraniales, series Disciflorae of subclass Polypetalae. However, all recent taxonomists,
including Cronquist (1981) and Thorne (1983) etc. treat it as an independent family belonging to order
Geraniales. Hutchinson placed the arborescent genus Averrhoa under a separate family Averrhoaceae.
Most of the taxonomists believe that Oxalidaceae have close affinities with Geraniaceae.
21.25 RUTACEAE CITRUS FAMILY OR RUE FAMILY


Distinguished by the presence of translucent, very clear dots in the leaves, Rutaceae are usually
shrubs or trees with aromatic oil glands; inflorescence cymose; outer stamens usually opposite the
petals; ovary deeply lobed and elevated on a disc.

Citrus, Aegle, Murraya, Ruta, Zanthoxylum, Limonia.

A family of about 150 genera and over 1500 species, Rutaceae are distributed in both tropical and
temperate regions, especially in Australia and South Africa. About 25 genera and over 80 species of
this family have so far been reported from India. Known throughout the world for its several juicy
citrus fruits, such as our present-day Oranges, Lemons, Grape-fruits, etc., Rutaceous members have
high percentage of vitamin C as well as several alkaloids. Some of its larger genera along with their
approximate number of reported species in bracket include Fagara (250), Zanthoxylum (200), Ruta
(60), Glycosmis (60), Eriostemon (32), Atalantia (18), Citrus (12), Murraya (12) and Aegle (3).

General Habit Shrubs or trees, and rarely herbs (Boenninghausenia albiflora, Ruta graveolens);
often xerophytic and aromatic or strong-smelling.
Root Branched tap roots.
Stem Erect, woody, branched; often with spines or thorns (Aegle, Citrus); usually aromatic.
Leaf Exstipulate; alternate (Citrus, Aegle) or opposite (Evodea); simple or compound; compound
leaves may be pinnately or palmately compound; sometimes reduced to spines (Citrus, Aegle); usu-
ally gland-dotted (Fig. 21.35).
Inflorescence Generally cymose; rarely raceme (Feronia); sometimes axillary (Citrus).
Flower Bracteate, bracteolate (Ruta) or ebracteolate, bisexual or rarely unisexual (Toddalia,
Zanthoxylum), actinomorphic or rarely zygomorphic (Dictamnus, Cusparia), pentamerous or rarely
tetramerous, hypogynous; cup-like nectariferous disc is usually present between ovary and stamens;
rarely trimerous (Triphasia trifoliata).
Calyx 4 to 5 sepals, free (Ruta) or united (Murraya, Fig. 21.35); valvate (Murraya) or imbricate
or quincuncial (Citrus medica); cup-like in Correa speciosa; rarely sepals absent in some species
of Zanthoxylum.
Corolla 4 to 5 petals, sometimes none; usually polypetalous; imbricate or rarely valvate; bell-shaped
(Correa); white, yellow or red; gamopetalous in Correa and Almeidea.
Androecium Usually 8 or 10 stamens; obdiplostemonous; anthers dithecous; introrse; rarely numer-
ous stamens (Citrus, Aegle); filaments usually free, but in Citrus filaments of numerous stamens
are united in several groups showing polyadelphous condition (Fig. 21.36); stamens of outer whorl
change into staminodes in Tetractomia.
Gynoecium Usually 4 to 5 carpels, rarely less or indefinite; carpels usually united or sometimes
free at base and united above by style; tetra- to pentalocular, 1 to 2 ovules in each locule, axile
placentation; ovary superior; rarely ovary unilocular with parietal placentation (Feronia).
Fruit and Seed Fruit hesperidium (Citrus), capsule (Flindersia), samara (Ptelea), drupe (Skimmia), or
rarely schizocarp, or follicle. Seeds without endosperm, with curved or straight embryo. Polyembryony,
with as many as 13 embryos (Citrus), is common.
Pollination and Dispersal Pollination entomophilous. Flies, bees, etc. are attracted because of coloured
corolla and nectar secreted by the disc. Dispersal of seeds takes place mainly by animals and also
by man.
General Floral Formula Br, ≈, , K4–5 or (4–5), C4–5, A8–10, G(4–5).
278 Plant Taxonomy
stigma stigma stigma

stamen
style
petal style
stamen
ovary
ovule
disc
sepal
thalamus
pedicel sepal
Flower Gynoecium ovary disc
pedicel thalamus
L.S. Flower
ovary wall
locule
ovule
placenta
T.S. Ovary
leaflet
gland
leaf
flower bud
stem
Flowering Branch
Floral Diagram
Fig. 21.35 Murraya paniculata (L.) Jacq.

• Fruits: Rutaceae provides some of the world’s most important, tasty and juicy citrus fruits,
which are the richest source of vitamin C. Some of them are Citrus aurantifolia (Lime,
Kaghzi Nimbu), C. aurantium (Sour orange, Khatta) C. limettioides (Sweet lime, Meetha
Nimbu), C. limon (Lemon, Galgal), C. maxima (Shaddock, Chakotra), C. medica (Citron,
stigma flower
stamens
style
ovary
petal
ovule sepal
disc thalamus
pedicel
L.S. Flower
gland
anther
lobe
leaf
filament stem
Flowering Branch
Stamens
ovary wall
locule
ovule
placenta
Floral Diagram T.S. Ovary
Fig. 21.36 Citrus medica L.
Bara Nimbu), C. paradisi (grapefruit), C. reticulata (Orange, Shantra) and C. sinensis (sweet
orange, Malta or Mussambi). Some other famous fruits of the family include Aegle marme-
los (Bael, Bilva), Evodia fraxinifolia (Kanukpa), Feronia limonia (Wood-apple, Kaith),
Fortunella japonica (Kumquat) and Glycosmis pentaphylla (Ban Nimbu).
• Condiments: Bark of Zanthoxylum alatum (Tejpat) is a famous condiment and is also
used to clean teeth and as a carminative and stomachic. Leaves of Z. limonella are used
280 Plant Taxonomy
as condiment and its fruits are digestive and appetizing. Fruits of Z. nitidum are used as
condiment and its roots are used in toothache. The leaves of Murraya koenigii are mixed
with a little turmeric to make curry powder in India. Leaves of Evodia lunuranthenoa are
also used as a condiment.
• Medicinal Value: (i) A poultice of bark of Acranychia laurifolia is applied on ulcers and
sores; (ii) fruit pulp of Aegle marmelos is a mild laxative whereas its roasted fruits are used
against diarrhoea and dysentry, and its root and stem bark in intermittent fever; (iii) oil
from the fruits of Atalantia monophylla is useful in paralysis and rheumatism; (iv) leaves
of Barosma betulina are used in curing kidney diseases; (v) bark of Cusparia febrifuga is
used in malaria; (vi) bark of Dictamnus albus is useful in nervous diseases and intermit-
tent fevers; (vii) ripe fruits of Feronia limonia are taken as a cardiac tonic; (viii) roots of
Glycosmis pentaphylla are used in fever; (ix) bark and roots of Murraya koenigii are used
in intestinal disorders; (x) seeds of Peganum harmala are given in asthma, neuralgia and
rheumatism; (xi) leaves of Pilocarpus pinnatifolius are used in kidney troubles; (xii) dried
leaves of Ruta graveolens are used as nerve stimulant; (xiii) root bark of Toddalia asiatica
is a potent antimalarial drug; (xiv) seeds of Zanthoxylum alatum are used in cholera; (xv)
roots of Z. nitidum are used in toothache.
• Ornamental Plants: Amongst the ornamental shrub and tree genera of Rutaceae include
Phellodendron, Ptelea, Choisya, Skimmia and Zanthoxylum. Perennial herbs of ornamental
value include Dictamnus albus and Ruta graveolens. The fragrant-flowered Murraya panicu-
lata is commonly grown for decorative purposes. Some other ornamentals include Fortunella
japonica, Glycosmis pentaphylla, Luvunga scandens and Poncirus trifoliata.
• Timber: The wood obtained from Choisya, Chloroxylon swietennia, Feronia limonia,
Glycosmis pentaphylla, Murraya paniculata and Zanthoxylum alatum is used for veneering,
making walking sticks, agricultural implements, etc.
• Oils: Essential oils, obtained from the leaves and fruit rind of various species of Citrus,
Feronia, Murraya, etc., are used in medicine and perfumery. Seeds of Citrus bergamia are
the source of famous ‘bergamot oil’.

Rutaceae belongs to order Geraniales, series Disciflorae and subclass Polypetalae according to
Bentham and Hooker, who divided it into four tribes viz. Aurantieae, Ruteae, Toddalieae and
Zanthoxyleae. Takhtajan (1969) and Thorne (1983) placed it under order Rutales whereas Cronquist
(1981) placed it under order Sapindales. Recently, Hickey and King (1988) divided Rutaceae into 5
subfamilies viz. Rutoideae, Dictyolomatoideae, Spathelioideae, Toddalioideae and Aurantioideae, and
stated that “Rutaceae is most closely related to Zygophyllaceae and Cneoraceae on the one hand and
to Meliaceae, Burseraceae and Simaroubaceae on the other, all included in the order Sapindales”.

1. Murraya paniculata (L.) Jacq. (syn. M. exotica; Fig. 21.35)
Stem: Herbaceous, lower portions woody, branched, erect, solid. Leaf: Alternate, petiolate, exstipu-
late, compound, unipinnate and imparipinnate, 3–7 leaflets; leaflets ovate to elliptical or lanceolate,
entire, obtuse; unicostate reticulate; gland-dotted. Inflorescence: Dichasial cyme. Flower: Bracteate,
pedicellate, complete, actinomorphic, hermaphrodite, pentamerous, hypogynous, white; a nectar-
secreting disc is present below the ovary. Calyx: 5 sepals, gamosepalous, valvate. Corolla: 5 petals,
free, imbricate. Androecium: 10 stamens, arranged in 2 whorls of 5 each, free, dithecous, basifixed,
introrse. Gynoecium: Bicarpellary, rarely 3 carpels, syncarpous; superior, bilocular, 1 to 2 ovules
in each locule, axile placentation; a hypogynous nectariferous disc is present; stigma bifid. Fruit:
Berry.
Floral Formula: Br, ≈, , K(5), C5, A5+5, G(2).
2. Citrus medica (Citron or Bara Nimbu; Fig. 21.36)
Stem: Same as in Murraya. Leaf: Exstipulate, leaf base swollen, simple, alternate, entire, ovate,
obtuse or acute. Inflorescence: Solitary axillary or axillary umbellate cyme. Flower: Almost same
as in Murraya. Calyx: 5 sepals, free or united, valvate. Corolla: Same as in Murraya. Androecium:
Numerous stamens, polyadelphous, attached round a disc, anthers dithecous, dorsifixed or basifixed,
introrse. Gynoecium: 5 to many carpels, syncarpous, superior, multilocular, one or more ovules in
each locule, axile placentation; a nectariferous disc is present. Fruit: Hesperidium.
Floral Formula: Br or Ebr, ≈, , K(5), C5, A(μ) polyadelphous, G(5 – μ).
21.26 MELIACEAE MAHOGANY FAMILY


Evergreen or deciduous trees or shrubs; leaves exstipulate, alternate, pinnately compound; inflores-
cence axillary, panicles or cymose; flowers bisexual, actinomorphic, hypogynous; sepals and petals
4–5; stamens 8–10, monadelphous; a disc is usually present between gynoecium and androecium.

Azadirachta, Melia, Swietenia, Toona, Walsura.

A family of about 50 genera and about 1400 species, Meliaceae is exclusively a tropical region
family of the world. About 20 genera and over 75 species of Meliaceae are reported from India.
Common Indian species include Azadirachta indica (Neem), Melia azedarach (Bakain) and Toona
ciliata (Toon) whereas Swietenia is a timber-yielding species of tropical America. Xylocarpus is a
mangrove genus of Meliaceae.

General Habit Trees or shrubs, evergreen (Appanomixis) or deciduous (Melia, Toona), contain hard
scented wood.
Stem Woody, solid, erect, branched, hard.
282 Plant Taxonomy
Leaf Alternate; pinnately compound, twice or thrice pinnate in

Melia, decompound, or rarely simple; exstipulate, do not possess
pellucid dots; leaflets usually with entire or serrate margin and
oblique at the base.
Inflorescence Axillary panicles or cymose.
Flower Bracteate (Fig. 21.37), bracteolate or ebracteolate; bisex-
ual, but rarely unisexual and polygamodioecious (Appanomixis);
actinomorphic, hypogynous; usually with an annular or a tubular
disc between ovary and stamens.
Calyx 4 to 5 sepals, usually basally connate or free; small;
imbricate but rarely valvate or contorted.
Corolla Usually 4 to 5 petals, rarely few or more than 5, free
or united; imbricate but rarely valvate; usually adnate to the
Fig. 21.37 Floral diagram of staminal tube.
Azadirachta indica Juss. Androecium Usually 8 to 10 stamens, rarely less than 8 or more
than 10; monadelphous, rarely free (Toona, Cedrela); filaments
unite to form a columnar tube; anthers dithecous, sessile, dehiscing longitudinally; a disc is usually
present between androecium and gynoecium.
Gynoecium Bi- to pentacarpellary, syncarpous, superior; 2 to 5 locules, 2 or rarely more (up to 12
in Swietenia) ovules in each locule; axile placentation; style short or ill-developed; stigma capitate
or discoid or lobed.
Fruit and Seed Fruit a berry, capsule, or rarely a drupe. Seeds often winged (Toona, Khaya), with
or without endosperm; rarely arillate (Appanomixis); wings absent in Azadirachta, Carapa and
Melia.
Pollination and Dispersal Pollination generally entomophilous; insects are attracted because of the
nectar secreted by the intrastaminal disc. Seeds are dispersed mainly by birds or other animals.
However, winged seeds are dispersed by wind.
General Floral Formula Br, ≈, , K(4–5), C4–5, A(8–10), G(2 – 5).
21.26.6 Economic lmportance

• Timber: Excellent timber trees of Meliaceae include Amoora wallichii, Azadirachta indica
(Neem tree or Margosa tree), Carapa granatum, Cedrela odorata, Chikrassia tabularis,
Chloroxylon swietenia, Dysoxylum malabaricum (white cedar), Khaya senegalensis (African
Mahogany), Melia azedarach (Bakain), Sandoricum koetjape, Soymida febrifuga, Swietenia
mahogani (True Mahogany), Toona ciliata syn. Cedrela toona (Toon wood), etc.
• Medicinal Value: (i) Seeds of Neem or Margosa tree (Azadirachta indica) are the source
of margosa oil, used in skin diseases; (ii) bark of A. indica is used in malarial fever while
its tender twigs are used as ‘datoon’ and are very effective in pyorrhea; (iii) leaf juice of
Melia azedarach is anthelmintic and its seeds are used in rheumatism; (iv) fruits of Aglaia
stamens staminal
stigma tube
staminal
tube
ovary petal
style
sepal
pedicel
disc A Flower
ovule
sepal
thalamus pedicel
L.S. Flower
flower
inflorescence
leaf
Fruits
stem
Flowering Branch
anther lobes
connective
Floral Diagram A part of Staminal Tube
Fig. 21.38 Melia azedarach L.
odorata are used in leprosy and inflammation; (v) bark of Carapa granatum are used in
dysentery and stomach troubles; (vi) wood oil of Dysoxylum malabaricum is used in eye
and ear diseases; (vii) bark of Soymida febrifuga is a bitter tonic and used in intermittent
fever; (viii) bark of Walsura piscidia is stimulant, expectorant and used in skin diseases.
284 Plant Taxonomy
• Minor Uses: (i) Ornamental plants of the family include Aglaia odorata, Melia composita,
Walsura trijuga, etc.; (ii) Cedar oil used in microscopy is obtained from Cedrela odorata;
(iii) flowers of Chukrasia yield a red-coloured dye.

Bentham and Hooker placed Meliaceae under order Geraniales of series Disciflorae. Meliaceae is
related to Rutaceae on one hand and to Burseraceae and Simarubaceae on the other. Bessey and
Engler also included Meliaceae under Geraniales and believed that it originated from Malvales.
However, Hallier was of the opinion that Meliaceae was derived from Rutaceae. Meliaceae has been
placed under order Meliales by Hutchinson while under Rutales by Takhtajan. Cronquist (1981) placed
Meliaceae under Sapindales while Thorne (1983) placed it under Rutales.

Melia azedarach (syn. Bakain, Fig. 21.38)
Habit: A big tree. Stem: Woody, erect, branched, solid. Leaf: Alternate, exstipulate, compound,
bipinnate and imparipinnate; leaf base pulvinus; pinnae ovate to lanceolate; serrate, unicostate
reticulate. Inflorescence: Axillary panicle cyme. Flower: Bracteate, bracteolate, pedicellate, complete,
actinomorphic, hermaphrodite, pentamerous, hypogynous; contains a nectariferous disc below ovary.
Calyx: 5 sepals, fused, valvate. Corolla: 5 petals, free, imbricate. Androecium: 10 stamens, mona-
delphous; filaments form a staminal tube having a ten-toothed apex; dithecous, basifixed, introrse.
Gynoecium: 5–8 or many carpels, syncarpous, superior, 5 to 8 or many-loculed, 1 or 2 ovules in
each locule, axile placentation; stigma lobed or capitate; a nectariferous disc is present below the
ovary. Fruit: Drupe.
Floral Formula: Br, Brl, ≈, , K(5), C5, A(10), G(5 – μ).
21.27 CELASTRALES
1. Leaves simple accept Ampelideae; often armed with spines or climbing by hooks or
tendrils.
2. Flowers actinomorphic and bisexual.
3. Stamens as many or lesser than that of petals, and only rarely twice as many as petals.
4. Stamens usually opposite the petals.
5. Disc present.
Bentham and Hooker included 4 orders (=families) under Celastrales. These are Ampelideae
(=Vitaceae), Celastrineae, Rhamneae and Stackhousieae.
Engler and Diels included Vitaceae and Rhamnaceae under Rhamnales and families Celastraceae
and Stackhousiaceae under suborder Celastrineae of order Sapindales.
Only Rhamnaceae and Vitaceae are treated in this text.
21.28 RHAMNACEAE BUCKTHORN FAMILY

Polypetalae, Disciflorae, Celastrales.

Shrubs or trees, often armed with spines or climbing by hooks or tendrils; leaves stipulate; flowers
perigynous; stamens opposite the petals; fruit drupe or capsule; seeds hard.

Zizyphus, Helinus, Berchemia, Ventilago.
21.28.4 Size and Distribution

Rhamnaceae comprises 58 genera and about 900 species. The plants of this family are cosmopolitan
in distribution. Some of the larger genera of the family along with their approximate number of
reported species are Rhamnus (150), Phylica (150), Zizyphus (100), Ceanothus (55), Frangula (50),
and Colletia (17).

General Habit Mostly trees or shrubs, often climbing by their hooks (Ventilago) or tendrils (Gouania,
Helinus) or twining stems (Ventilago, Berchemia); some thorny (Colletia, Condalia); rarely herbs.
Leaf Simple, stipulate, alternate or rarely opposite; stipules modify into spines in Zizyphus; usually
pinninerved but palminerved in Zizyphus.
Inflorescence Axillary corymbs or cymose; solitary axillary in Microrhamnus.
Flower Bracteate, bisexual or rarely unisexual, actinomorphic, sometimes apetalous (Colletia),
pentamerous or tetramerous; perigynous or epigynous; usually small; hypogynous in Berchemia
scandens.
Calyx 4 or 5 sepals, free or basally connate, valvate.
Corolla 4 or 5 petals, free, usually concave and often clawed at
the base, small, valvate; sometimes absent (Colletia); petals often
inserted on the calyx tube to form a hypanthium.
Androecium Stamens as many as petals, opposite the petals
(Fig. 21.39) and enclosed by them, inserted on the hypanthium or
develop on the margin of intrastaminal disc; anthers dithecous;
Gynoecium Carpels 2–4, syncarpous or free, bi- to tetralocular,
ovary superior or half-inferior, one or 2 ovules in each locule, basal
placentation; style simple or divided; stigma simple or bilobed.
Fruit and Seed Capsule (Phylica, Pomaderis), drupe (Rhamnus, Fig. 21.39 Floral diagram
Zizyphus) or samaroid (Ventilago), or nut. Seeds hard, large, with of Zizyphus jujuba.
straight embryo and little or no endosperm.
286 Plant Taxonomy
Pollination and Dispersal Pollination mostly entomophilous, and majority of the dry fruits are dis-
persed by wind.
General Floral Formula Br, ≈, , K4–5, C4–5, A4–5, G(2 – 4).

• Ceanothus (Californian Lilacs) is the most commonly grown genus for decorative purposes
because of its beautiful blue, pink or white flowers. Some other ornamental plants include
species of Berchemia, Colletia, Discaria, Paliurus, etc.
• Zizyphus mauritiana (Baer of India), Z. nummularia (Jhar Ber), Z. jujuba, Z. vulgaris,
Sageretia oppositifolia, Hovenia dulcis (Japanese Raisin-tree) are well known for their edible
fruits.
• Green and yellow-coloured dyes of commercial importance are obtained from various spe-
cies of Rhamnus. A red dye is obtained from root bark of Ventilago maderaspatana.
• Cascara sagarda bark, a purgative, is obtained from Rhamnus purshiana.
• Gun powder is prepared from the wood of Zizyphus mauritiana.
• Ceanothus americanus (New Jersey Tea) leaves are used as a substitute of tea.
• Valuable timber is obtained from Frangula alnus, Maeosopis emini and Zizyphus chloroxy-
lon. Wood of Gouania domingensis is used for making chew-sticks.

Bentham and Hooker placed Rhamnaceae under order Celastrales of series Disciflorae. But majority
of the other taxonomists place it under order Rhamnales (Takhtajan, 1969; Cronquist, 1981; Thorne,
1983). Rhamnaceae shows affinity with Vitaceae from the standpoint of floral construction. However,
it differs from Vitaceae by its small petals, receptacle, endocarp and always simple leaves. Some
believe Rhamnaceae to be close to Celastraceae but the main distinction between the two is the
presence of antipetalous stamens in Rhamnaceae.
21.29 VITACEAE GRAPE FAMILY

Polypetalae, Disciflorae, Celastrales.

Climbing habit; terminal buds of plants develop into tendrils; inflorescence opposite the leaves;
stamens usually opposite petals; ovary bilocular; axile placentation; fruit berry.

Ampelocissus, Cayratia, Cissus, Leea and Vitis.

A small family of only 12 genera and 700 species, Vitaceae are mostly distributed in tropical and
subtropical regions of the world. 8 genera and about 70 species of Vitaceae have been reported from
India, mostly growing in plains. Most important and universally known plant of the family is Vitis
vinifera (Grape vine). Some larger genera of Vitaceae along with their approximately reported species
are Cissus (350), Ampelocissus (95), Tetrastigma (90), Vitis (60 to 70) and Parthenocissus (15).

General Habit Most genera are vines or climbing shrubs; rarely erect shrubs, or small tree
(Carameriana); species of Leea are erect herbs, shrubs or even trees.
Stem Hard, woody, with nodes often swollen or jointed; tendrils are leaf-opposed and considered
as the metamorphosed apical part of the sympodial stem by Eichler. (Tendril is a negatively geotro-
pic organ. The tips or ends of tendrils expand to form sticky and mucilaginous, ball-like structures,
which help them to adhere firmly to the support.)
Leaf Alternate, but lower leaves are sometimes opposite; simple or compound (pinnately or pal-
mately); stipules petiolar or absent; with pellucid punctate dots.
Inflorescence Flowers borne opposite the leaves in racemose, paniculate or cymose manner.
Flower Bracteate, usually bracteolate, actinomorphic, bisexual or unisexual; 4 to 5-merous, hypogy-
nous; minute; peduncle is flat and ribbon-like in Pterisanthes.
Calyx 4 to 5 sepals, rarely 3 to 7, fused; very small, cupular, valvate.
Corolla 4 to 5 petals, free or fused; caducous and fall easily when the flowers open; valvate; in
Vitis petals are free near base and united by the apices.
Androecium 4 to 5 stamens, opposite the petals (Fig. 21.40); arise from the base of an intrastaminal
disc which is well-developed, annular or lobed; anthers free or connate, dithecous, introrse; dehis-
cence longitudinal; in Leea, the filaments unite and form a 5-lobed disc.
Gynoecium 2 to 8 carpels, syncarpous, superior, 2 to 8-loculed, 1 or 2 ovules in each locule, axile
placentation; style one and short; stigma discoid or capitate.
Fruit and Seed Fruit berry, often with watery juice. Seeds with copious endosperm and straight
embryo.
Pollination and Dispersal Pollination is entomophilous and dispersal takes place usually by birds,
animals or even man.
General Floral Formula Br, Brl, ≈, , K(4–5), C4–5, A4–5, G(2 – 8).

Vitaceae are important for the table grapes, wine grapes, raisins, grape juice, and for several orna-
mental and medicinal purposes.
• Fruits: Principal economic plant is Vitis vinifera (Grape vine, Fig. 21.40), whose several
varieties are grown for their edible fruit, the source of raisins, wines, sultanas, and currants.
Other species of Vitis known for their less palatable edible fruits include V. aestivalis and
V. labrusca. Fruits of Leea crispa and L. macrophylla are also edible.
288 Plant Taxonomy
stigma
style
ovary
T.S. Ovary T.S. Ovary
(bilocular) (trilocular)
stamens
L.S. Ovary disc
stigma
style Flower
(after shedding
Stigma with petals)
Style
leaf
stem Floral Diagram
inflorescence
petals
Fruits
A Floral Bud Flowering Branch
Fig. 21.40 Vitis vinifera L.
• Ornamental Vines: Parthenocissus tricuspidata (Japanese creeper), P. quinquefolia

(Virginia creeper) and several species of Ampelopsis, Cissus, Rhoicissus, etc. are cultivated
as ornamental vines on walls.
• Medicinal Value: (i) Leaves and roots of Cayratia carnosa are used in high fever; (ii) tubers
of Cissus adnata are used as blood purifier; (iii) the essential oil obtained from the plant
of Leea aequata are used against tuberculosis; (iv) roots of L. indica are used in diarrhoea;
(v) tubers of L. macrophylla are used in ringworm disease.
• Minor Uses: (i) Indian lac insect is grown on Leea crispa and L. edgeworthi; (ii) leaves of
several species of Parthenocissus are used as fodder; (iii) a strong cordage fibre is obtained
from the stems of Cissus adnata and C. quadrangularis.

Bentham and Hooker included Vitaceae (= Ampelideae) under order Celastrales of series Disciflorae
whereas majority of the other authors (Hutchinson, Takhtajan, Cronquist, Thorne, etc.) place it under
order Rhamnales. Vitaceae is closely allied to Rhamnaceae due to their similar floral structure.
However, it differs from Rhamnaceae by its climbing habit, berried fruit, small embryo and albu-
minous seeds.
21.30 SAPINDALES
1. Usually woody shrubs or trees.
2. Leaves usually compound and exstipulate.
3. Flowers small, often irregular, pentamerous and hypogynous.
4. Stamens definite, usually inserted on a prominent receptacular or extrastaminal disc.
5. Ovary tricarpellary, syncarpous, superior, usually trilocular.
6. Ovules pendulous with the dorsal raphe and micropyle upward or erect with ventral raphe
and micropyle downward.
Sapindales includes Sapindaceae, Anacardiaceae and Sabiaceae according to Bentham and Hooker.
Engler and Diels treated Sapindales as composed of 11 suborders and 23 families, of which some
major ones are Buxaceae, Anacardiaceae, Celastraceae, Salvadoraceae, Sapindaceae and Sabiaceae.
Only Sapindaceae and Anacardiaceae are treated in this text.
21.31 SAPINDACEAE SOAPBERRY FAMILY

Polypetalae, Disciflorae, Sapindales.

Usually trees or shrubs with pinnately compound leaves; flowers small, pentamerous and hypogy-
nous; sepals imbricate; petals glandular; an extra-staminal disc is usually present; ovary tricarpellary,
syncarpous.

Litchi, Sapindus, Acer, Aesculus, Dodonaea and Schleichera.
290 Plant Taxonomy
21.31.4 Size, Distribution and General lnformation

A family of about 155 genera and over 2220 species, Sapindaceae are distributed mainly in tropi-
cal and subtropical regions of the world. About 25 genera and 75 species of this family have been
reported from India, mainly from tropical Eastern Himalayas and Western Peninsular India. Sapindus
saponaria (soapberry tree) provides the common name ‘Soapberry family’ to this family. Litchi chin-
ensis (Litchi), Acer saccharinum (Sugar Maple), Paullinia (Guarana) and Cardiospermum (Balloon
vine) are other famous genera of the family.

General Habit Trees or shrubs, rarely herbs, sometimes tendril-bearing vines (Cardiospermum) or
lianas (Serjania, Paullinia); tendrils being the flowerless branches of inflorescence.
Leaves Usually alternate, pinnately compound and exstipulate; very rarely opposite (Acer,
Fig. 21.41A); simple (Cardiospermum); small and deciduous stipules present in climbing species;
often contain resin or latex cells.
Inflorescence Flowers minute and usually arranged in racemose to paniculate unilateral cymes or
a thyrsus; plants commonly polygamodioecious (i.e. bear apparently bisexual but functionally uni-
sexual flowers); metamorphosed sterile branches of inflorescence become coiled like watch-spring
and appear like tendrils.
Flower Minute, bracteate or ebracteate, bracteolate (Acer), bisexual or unisexual, actinomorphic
(Litchi, Aphania, Sapindus) or obliquely zygomorphic (Cardiospermum, Erioglossum), pentamerous
or rarely tetramerous, hypogynous.
Calyx 5 sepals, poly- or gamosepalous, imbricate or rarely valvate; sometimes only 4 sepals due
to the fusion of 3rd and 5th sepals.
Corolla 5 petals, sometimes 3, or even absent (Schleichera), polypetalous, valvate (Acer) or imbri-
cate (Sapindus); petals equal or unequal-sized and often bear scale-like or hairy appendage at the
base.
Androecium Stamens 8 (Acer) or 10 (Sapindus, Fig. 21.41E), free and inserted inside a prominent
receptacular or extrastaminal disc; only 5 stamens in Turpinia, and numerous stamens in Deinbollia;
filaments often hairy; zygomorphic flowers often have eccentric androecium; anthers 2-celled, bas-
ifixed or versatile and introrse. In between petals and stamens is often present an annular disc.
Gynoecium Tricarpellary, syncarpous; ovary superior; trilocular with one or two or rarely more
ascending ovules in each locule, axile placentation; style simple or divided; rarely the placentation
is parietal.
Fruit and Seed Fruits may be dry or fleshy and highly variable from capsule, berry, nut, drupe,
samara, or schizocarp in different genera. Seeds are non-endospermic with a curved embryo; often
arillate.
Pollination and Dispersal Pollination is by insects such as flies, bees, etc. Dispersal takes place by
wind, birds or other animals.
General Floral Formula Br or Ebr, ≈, , K5, C5 or zero, A8 – 10, G(3).
inflorescence
C
A Flower
Fruits (viewed from top)
leaf
stem
A
Flowering
Branch
E
D
Floral Diagram Floral Diagram
Fig. 21.41 A–D: Acer oblongum Wall, ex D, C.; E: Floral diagram of Sapindus.
292 Plant Taxonomy

The economic importance of Sapindaceae lies in the fact that it provides several edible fruits, e.g.
Litchi; saponin, guarana, akee, plus some ornamental plants.
• Delicious edible fruits are obtained from Litchi chinensis (Litchi), Blighia sapida (Akee),
Euphoria longana (Anshphal), Melicocca bijuga and Nephelium lappaceum (Rambutan).
The edible part of Litchi is sweet pulpy aril which surrounds the seed.
• Acer hippocastanum (Horse chestnut tree) fruits are given to horses as well as used for
manufacture of starch; A. indica is a timber-yielding plant and its fruits and seeds are used
in rheumatic complaints; A. saccharinum (Sugar maple) is the source of maple sugar and
timber.
• Dodonaea viscosa (Villayati Mehndi), an evergreen shrub, is a popular ornamental hedge
plant.
• Nephelium lappaceum (Rambutan of Malaya) fruits are edible and the fat obtained from its
seeds is used in making candles and soap.
• Paullinia cupana (Guarana) seeds are used like cacao in Brazil to make a drink having
high caffeine content.
• Sapindus mukorossi (Ritha or Soapnut tree) and S. saponaria (Soapberry tree) fruits contain
saponin. On being moistened, saponin forms a lather and used as a soap. Ritha is used for
washing hairs and clothes. Saponin is used in the preparation of soaps, tooth pastes, sham-
poos as well as in some insecticides.
• Schleichera oleosa yields edible fruit and this tree is used for rearing lac insects. It also
provides a good timber and the oil obtained from its seeds is used for cooking.

Sapindaceae was placed under order Sapindales (after Celastrales and before Rosales) of series
Disciflorae by Bentham and Hooker. Order Sapindales was placed between Rutales and Rhamnales
by Rendle (1925), Geraniales and Rhamnales by Lawrence (1951), and between Polygalales and
Geraniales by Cronquist (1981). Thorne (1983) discussed Sapindaceae under order Rutales, suborder
Sapindineae.
Sapindaceae is related to Anacardiaceae in habit and general floral structure. However, the flow-
ers are mostly irregular in Sapindaceae and regular in Anacardiaceae. In the characters, such as the
presence of one ovule in each locule, tricarpellary ovary, often unisexual flowers and arillate seeds,
family Sapindaceae resembles Euphorbiaceae.
Sapindales are generally thought to be evolved parallel to Rutales and Meliales.
Bentham and Hooker divided Sapindaceae into five subfamilies (Sapindeae, Acerineae, Dodoneae,
Meliantheae and Staphyleae). However, the recent trend is to divide it into only two subfamilies viz.
Sapindoideae and Dodonaeideae.
21.32 ANACARDIACEAE CASHEW FAMILY

Polypetalae, Disciflorae, Sapindales.

Trees or shrubs; pentamerous flowers; presence of intrastaminal disc; presence of resin ducts; uni-
locular ovary; fruit usually drupe.

Mangifera, Rhus, Anacardium, Spondias, Buchanania and Lannea.

A family of about 80 genera and over 600 species (Jones and Luchsinger, 1987), Anacardiaceae are
mostly tropical in distribution but found also in temperate regions of Mediterranean, America and
eastern Asia. 23 genera and over 115 species of Anacardiaceae have been reported from India. Some
larger genera of Anacardiaceae, along with their common names and approximately reported species
in parenthesis, include Rhus (Sumac or poison ivy, 250), Semecarpus (Dhobis nut, 50), Mangifera
(Mango, 40), Schinus (Pepper tree, 30), Anacardium (Cashew nut, 15), Pistacia (Pistachio nut,
10), Spondias (Hog plum, 10) and Cotinus (Smoke tree, 3). Mangifera indica (mango) is the most
familiar plant of the family.

General Habit Usually trees and shrubs, rarely climbers or woody vines; containing resin ducts with
plenty of gum and acrid juice.
Leaves Usually alternate (opposite in Dobinea, Bouea), simple or pinnately compound; exstipulate
or stipules obscure.
Inflorescence Axillary or terminal panicles.
Flower Bracteate, bisexual or sometimes unisexual, pentamerous, actinomorphic; hypogynous, rarely
perigynous (Holigarna) or epigynous; small and numerous.
Calyx Usually 5 sepals, sometimes 3 to 7; free or basally connate or adnate to gynoecium, or vari-
ously divided; imbricate or quincuncial.
Corolla Usually 5 petals, sometimes 3 to 7; or even absent (Pistacia); free or rarely fused; imbricate
or rarely valvate.
Disc A cupular, extrastaminal or intrastaminal, nectariferous disc is usually present; rarely the disc
is absent (Pistacia).
294 Plant Taxonomy
longest longest
stamen stamen stamen
petal
lateral style short

stamens
ovary
short
stamens
A Flower sepal
Flower (with removed

L.S. Flower corolla)
flower
fleshy
pseudocarp fruit
T.S. Ovary
true fruit (nut)

leaf
Fruit
stem
Flowering Branch
Floral Diagram
Fig. 21.42 Anacardium occidentale L.
Androecium 10 stamens (or stamens double the number of petals), in 2 whorls; usually fewer stamens
(1 stamen and 6–9 staminodes or ill-developed stamens in Anacardium, Fig. 21.42; 1 stamen and
4 staminodes in Mangifera, Fig. 21.43; 5 stamens in one whorl in Rhus, Fig. 21.44); androecium
fertile
reduced stamen
stamen
petal
sepal
pedicel
A Flower
stigma
style
ovary
petal
ovule disc
sepal
pedicel
L.S. Flower Floral Diagram
mesocarp
endocarp
inflorescence seed
epicarp
L.S. Fruit
leaf
stem
Flowering Branch
Fig. 21.43 Mangifera indica L.

296 Plant Taxonomy
usually arise from beneath a disc; filaments free or rarely

basally connate (Anacardium); anthers basifixed or dorsi-
fixed, introrse.
Gynoecium A compound pistil of 1 to 5 fused carpels to
form a unilocular or multilocular superior ovary, having a
solitary ovule in only one of the ovary chamber; sometimes
3 carpels; styles 1–5; each with a capitate stigma; only one
carpel in Anacardium and Mangifera; in Drimycarpus the
ovary is inferior; placentation basically axile.
Fruit and Seed Fruit usually a drupe (Mangifera) with resin-
uous mesocarp; in Anacardium the fruit is a nut and remains
situated on a pyriform fleshy mass developed by the elonga-
tion of disc and top of pedicel. Seed contains curved embryo
Fig. 21.44 Floral diagram with very little or no endosperm.
of Rhus parviflora. General Floral Formula Br, ≈, , K5, C5, A10, G(1 – 5).

Anacardiaceae is important for the mango, cashew, pistachio, resins, oils, tannic acids, chironji,
lacquers, several ornamentals, as well as for causing dermatitis in humans.
• Mangifera indica (mango) is universally known for its delightful and delicious fruit as well
as for providing us pickles and timber.
• Anacardium occidentale (Cashew-nut) is famous for its edible nuts (seeds). The fruit and
seed coat of the cashew contain a poison. However, its seeds are edible and highly priced.
• Buchanania lanzan (Chironji) seeds are commonly used in sweet-meats in India. Its bark
is used for tanning.
• Harpephyllum caffrum (kaffir plum) is famous for its edible fleshy fruits.
• Holigarna arnottiana provides useful timber for match industry and packing cases.
• Lannea coromandelica bark is used for tanning and provide a gum used in calico
printing.
• Parishia insignis provides useful timber for railway sleepers.
• Pistacia vera (pistachio-nuts) is the source of famous edible nuts used in sweet-meats, ice-
creams, etc. P. lentiscus (mastic tree) yields a mastic resin used in chewing gums, alcoholic
beverages, etc.
• Rhus vernicifera (Lacquer tree) is grown for its milky poisonous latex. Contact with this plant
may cause dermatitis in humans. Tannic acids are obtained from R. coriaria. R. parviflora
(Tatri) fruits are used as tamarinds.
• Schinopsis lorentzii is used in leather-tanning industry.
• Semecarpus anacardium (Dhobis-nut) fruits provide a black ink used for dyeing textiles.
• Spondias axillaris and S. pinnata (Hog plum) fruits are edible and made into pickles.
• Toxicodendron vernicifera provides us resins and oils.
• Some ornamental plants of Anacardiaceae include Cotinus coggygria, Rhus typhina and
Spondias pinnata.

Bentham and Hooker, Engler and Prantl, Cronquist and several other taxonomists placed Anacardiaceae
under Sapindales while Takhtajan (1969) and Thorne (1983) placed it under order Rutales.
Majority of the phylogenists trace the affinity of Anacardiaceae with Sapindaceae. The family
is, however, related closely to Aceraceae, Hippocastanaceae, Sapindaceae, Meliaceae, Rutaceae and
Burseraceae. The recent trend is to divide Anacardiaceae into 5 tribes viz. Anacardieae, Spondiadeae,
Rhoeae, Semecarpeae and Dobineeae (Hickey and King, 1988).
21.33 ROSALES
1. Flowers bisexual, generally cyclic and typically pentamerous.
2. Flowers hypogynous to epigynous with common perigyny.
3. Stamens commonly arranged in many whorls.
4. Gynoecium apocarpous to syncarpous but the styles general distinct.
5. Ovary generally inferior, less-commonly half-inferior or superior.
6. Thalamus discoid or concave or flask-shaped.
Bentham and Hooker included 9 orders (=families) under Rosales, of which the major ones are
Leguminosae, Rosaceae and Saxifragaceae. Order Rosales was considered by Engler and Diels to be
composed of 17 families, chief amongst which are Crassulaceae, Saxifragaceae, Hamamelidaceae,
Rosaceae and Leguminosae.
Only Leguminosae, Rosaceae and Saxifragaceae are treated in this text.
21.34 LEGUMINOSAE LEGUME FAMILY

Polypetalae, Calyciflorae, Rosales.
Traditional approach is to treat all legumes as one large, somewhat heterogenous family, the
Leguminosae under order Rosales. Bentham and Hooker followed this approach and divided family
Leguminosae into three subfamilies (Papilionaceae, Caesalpinieae and Mimoseae). However, majority
of the recent taxonomists (Hutchinson, 1973; Takhtajan, 1980; Cronquist, 1981, etc.) treat all these
three as three separate families (Mimosaceae, Caesalpiniaceae and Fabaceae or Papilionaceae) under
order Fabales (Takhtajan, 1980; Cronquist, 1981) or Leguminales (Hutchinson, 1969). The traditional
298 Plant Taxonomy
approach of Bentham and Hooker is followed in this book, and the three have been treated as
subfamilies under the names Mimosoideae, Caesalpinioideae and Papilionoideae. However, some
generalised characters of Leguminosae are given in the following paragraph:
Leguminosae, represented by about 600 genera and over 12000 species, is the third largest
family of the flowering plants. Plants are herbs, shrubs, trees, or vines. Leaves simple or pinnately
compound, often trifoliate, rarely palmately compound, alternate or opposite, usually stipulate; at
the base of petiole or petiolule is often present a pulvinus; rarely leaves are reduced to scales.
Inflorescence racemose raceme, panicle, spike, head, or flowers solitary. Flower bisexual, actinomor-
phic (Mimosoideae) or zygomorphic (Caesalpinioideae and Papilionoideae, Fig. 21.45), hypogynous
or rarely perigynous. Calyx of usually 5 connate sepals, imbricate or rarely valvate. Corolla of 5
petals, rarely less, polypetalous or 2 anterior petals fuse along their lower margins to form keel,
equal or unequal, imbricate or rarely valvate. Androecium of 10 stamens (numerous in Mimosoideae),
rarely less than 10, polyandrous, or monadelphous, or diadelphous, usually longitudinally dehiscent.
Gynoecium monocarpellary, ovary superior or rarely semi-inferior, unilocular, marginal placentation,
style and stigma one and simple. Fruit usually a legume, or lomentum. Seeds usually with a shining
leathery testa; endosperm very scanty or even absent.
style
uppermost
petal
stamen
staminal tube
stamen
corolla
style
calyx
Mimosoideae Caesalpinioideae
standard wing
keel (2 petals) sepal
Papilionoideae
Fig. 21.45 Flowers of Leguminosae.

21.34.2 Key to Subfamilies

Flowers actinomorphic; perianth aestivation valvate or rarely imbricate; stamens 10 or usually
more. .................................................................................................................................... Mimosoideae
Flowers zygomorphic; perianth aestivation usually imbricate; stamens between 5–10.
Corolla aestivation ascending-imbricate; posterior petal innermost; petals 5 and polypetalous; stamens
polyandrous or monadelphous. .....................................................................................Caesalpinioideae
Corolla aestivation descending imbricate; posterior petal outermost; petals 5 but anterior petals basally
fused; stamens monadelphous or diadelphous. ...............................................................Papilionoideae.
21.35 SUBFAMILY MIMOSOIDEAE OR MIMOSEAE MIMOSA FAMILY

Polypetalae, Calyciflorae, Rosales, Leguminosae.

Trees or shrubs, rarely herbs; leaves bipinnately compound; stipules usually represented by a pair
of thorns; flowers pentamerous, actinomorphic; stamens 10 to many; filaments coloured and often
much longer than corolla.

Acacia, Albizzia, Leucaena, Mimosa.

Represented by about 56 genera and 2800 species, Mimosoideae are predominantly tropical and
subtropical in distribution and almost absent in the colder regions of the world. About 15 genera and
75 species of this subfamily have so far been reported from India. Some of the larger genera along
with their number of approximately reported species include Acacia (750), Mimosa (450), Albizzia
(100), Desmanthus (40), Prosopis (40), Schrankia (30), and Neptunia (11). Mimosa pudica (sensitive
plant) is the best known plant of this subfamily.

General Habit Mostly medium-sized trees or shrubs (Albizzia, Acacia, Dicrostachys, Parkia, Xylia,
etc.), few are herbs (Neptunia oleracea) or woody and tendril-bearing climbers (Entada).
Leaves Mostly bipinnate; some species of Acacia appear to possess simple leaves, which are
actually phyllodes or leaf-like petioles having no blade; stipulate, stipules often modify into thorns
(Acacia); sensitive or possess sleep-movement character in Mimosa pudica (Fig. 21.46) and Neptunia
oleracea.
Inflorescence Flowers often grouped in tight clusters, or racemose head or capitulum, sometimes
spicate or racemose (Prosopis).
300 Plant Taxonomy
ovary wall ovule

stigma
anther
lobe
locule
filament
style T.S. Ovary
petal stigma
style
Fruit
ovary
calyx
A Flower
inflorescence
Gynoecium
leaflet
leaf
fruit
stem
Floral Diagram
Flowering Branch
Fig. 21.46 Mimosa pudica L.
Flower Bracteate, actinomorphic, bisexual, pentamerous, sometimes tetra-, tri- or hexamerous; hyp-
ogynous, rarely perigynous.
Calyx 5 sepals, fused into a 5-lobed tube; valvate or rarely imbricate (Parkia); sometimes 4 diago-
nally placed sepals (Mimosa).
Corolla 5 petals, fused (Acacia, Albizzia) or free (Parkia, Mimosa, Dicrostachys); valvate; 4 petals
in Mimosa (Fig. 21.46).
Androecium 4 to numerous stamens, show much variation in number and cohesion; extend beyond
the petals and provide a fluffy soft appearance to the floral cluster; stamens are 4 or as many as
petals (Mimosa, Fig. 21.46), or double the number of petals and diplostemonous (Prosopis), or
stigma
style
leaflet
inflorescence
ovary
Fruit stipule
Gynoecium
leaf
stem
Flowering Branch stigma
stamen
petal
style
sepal
ovary
L.S. Flower
ovary wall
locule
ovule
Floral Diagram T.S. Ovary
Fig. 21.47 Acacia nilotica L.
numerous and free (Acacia, Fig. 21.47), or numerous and monadelphous (Albizzia); dithecous, bas-
ifixed, introrse.
Gynoecium Monocarpellary; ovary superior, unilocular; ovules numerous, marginal placentation;
style and stigma simple.
Fruit A legume, sometimes indehiscent, lomentum in Acacia.
Seed Non-endospermic or with scanty endosperm; embryo straight.
General Floral Formula Br, ≈, , K(5), C5 or (5), A4 or 10 or μ, G1.
302 Plant Taxonomy

• Ornamental Plants: Acacia arabica syn. A. nilotica (Babool or Kikar), A. concinna
(Rassaul), A. farnesiana (Vilayati kikar), A. melanoxylon (Australian Acacia), Albizzia
lebbek (Siris), Leucaena leucocephala (Subabul), Mimosa pudica (sensitive plant) and
Pithecellobium dulce are the commonly grown ornamental plants.
• Catechu of Commerce or Katha: Katha, used for colouring khaki clothes and taken with
betel leaves, is obtained from the heartwood of Acacia catechu.
• Gum Arabic: True gum arabic, used in medicine, confectionery and textile industry, is
obtained from Acacia senegal and A. stenocarpa.
• Tannins: Bark of several species of Acacia (A. catechu, A. decurrens, A. leucophloea, A.
pennata, etc.) is rich in tannins and is used in tanning of animal skins.
• Timber: Valuable timber for furniture, cabinet work, agricultural implements and other
similar purposes is obtained from various species of Acacia (A. catechu, A. ferriginea, A. leu-
cophloea, A. nilotica), Albizzia (A. amara, A. chinensis, A. lebbek, A. procera), Adenanthera
pavoniana and Xylia xylocarpa.
• Edible Products: Beans of Entada phaseoloides, leaves of Neptunia oleracea and seeds
of Pithecellobium dulce are edible. Fruits and seeds of Prosopis cineraria and Samanea
saman are used as fodder.
• Miscellaneous Uses: (i) Acacia concinna (Shikakai) pods are used for washing hairs;
(ii) A. decurrens and Xylia xylocarpa wood pulp is used for making wrapping paper;
(iii) A. farnesiana flowers provide us a delicate perfume; (iv) A. nilotica plants are used
as host for lac insects; (v) A. sinuata pods are used for manufacturing of shampoos;
(vi) Adenanthera pavoniana seeds are used as weights by jewellers; (vii) Entada phase-
oloides seeds are used for washing hairs.
21.35.7 Description of a Common Representative

1. Acacia nilotica (L.) Del. syn. A. arabica Willd (Kikar or Babool; Fig. 21.47)
Habit: Medium-sized tree. Leaf: Alternate; stipulate, stipules modify into long thorns; bipinnate
and paripinnate. Inflorescence: Axillary cymose head. Flower: Bracteate, complete, actinomorphic,
hermaphrodite, hypogynous, pentamerous, numerous, small. Calyx: 4 or 5 sepals, fused basally,
valvate. Corolla: 4 or 5 petals, free or connate at the base, valvate. Androecium: Stamens numer-
ous, polyandrous, filaments long; anthers dithecous, basifixed, introrse. Gynoecium: Monocarpellary,
ovary superior; unilocular; many ovules, marginal placentation. Fruit: Lomentum.
Floral Formula: Br, ≈, , K(4–5), C4–5 or (4–5), Aμ, G1.
21.36 SUBFAMILY CAESALPINIOIDEAE OR CAESALPINIEAE CASSIA FAMILY


Trees, shrubs, or herbs; flowers zygomorphic; corolla aestivation ascending-imbricate; posterior petal
innermost; petals 5 and polypetalous; stamens usually 10, rarely numerous.

Bauhinia, Caesalpinia, Cassia, Delonix, Parkinsonia, Saraca, Tamarindus.

Represented by about 180 genera and 3000 species, members of subfamily Caesalpinioideae are
distributed mostly in tropical and subtropical countries. About 23 genera and 85 species of this
subfamily have been reported from India. Some of the largely represented genera along with
their number of approximately reported species include Cassia (500–600), Bauhinia (300), Senna
(250), Chamaecrista (250), Caesalpinia (100), Gleditsia (11) and Cercis (7) as stated by Jones and
Luchsinger (1987) and Hickey and King (1988).

General Habit Mostly trees (Tamarindus) and shrubs (Parkinsonia); rarely herbs (Cassia tora),
or prickly climbers (Caesalpinia sepiaria); Cassia comprises of all the three habits i.e. herbs (C.
pumila), shrubs (C. timoriensis) and tall trees (C. javanica).
Leaves Usually alternate, pinnate or bipinnately compound; rarely simple (Bauhinia); stipules some-
times modify into spines (Parkinsonia).
Inflorescence Usually raceme, or spikes, rarely corymb; or flowers solitary.
Flower Bisexual, zygomorphic, pentamerous, hypogynous or perigynous; rarely unisexual
(Gleditsia).
Calyx Usually 5 sepals, free or basally connate, imbricate or valvate; odd sepal anterior; sometimes
2 posterior sepals fuse (Tamarindus indica, Fig. 21.48A); sepals petaloid in Saraca indica.
Corolla 5 petals, polypetalous, ascending-imbricate aestivation; showing posterior petal inner-
most; usually the petals are unequal-sized; petals greatly reduced in Dialium; petals absent in
Ceratonia.
Androecium Usually 10 free stamens, arranged in 2 alternate whorls of 5 each (diplostemonous);
rarely numerous and united below; often a few stamens reduced to staminodes (e.g., number of perfect
stamens is only 3 in Tamarindus; 3 to 5 in Bauhinia, Fig. 21.48B; and 5 to 7 in Cassia, Fig. 21.49;
while their all remaining stamens are reduced to staminodes); filaments often unequal; anthers dith-
ecous, introrse and dehisce by apical pores (Cassia) or longitudinal slits (Delonix, Parkinsonia).
Gynoecium Monocarpellary, superior; unilocular ovary, with two alternating rows of anatropous or
campylotropous ovules on marginal placentation; style single, ending into a simple stigma.
Fruit and Seeds Fruit legume, sometimes indehiscent; often winged. Seeds endospermic or non-
endospermic.
General Floral Formula , , K5 or (5), C5, A10 or (μ), G1.
304 Plant Taxonomy
A B
Fig. 21.48 A: Floral diagram of Tamarindus indica; B: Floral diagram of Bauhinia variegata.

Subfamily Caesalpinioideae is important as a source of several ornamentals, drugs, dyes and
timber.
• Ornamental Plants: Amherstia nobilis (Noble Amherstia), Bauhinia acuminata (Safed
Kachnar), B. corymbosa (white-flowers), B. galpini (scarlet flowers), B. monandra (hedge
plant), B. purpurea (purple-flowered Kachnar or Orchid tree), B. racemosa (pink-flowered
Kachnar), B. tomentosa (yellow-flowered), B. variegata (variegated-flowered Kachnar),
Caesalpinia pulcherrima (orange or yellow-flowered Poinciana), Cassia alata, C. fistula, C.
javanica, C. marginata, C. roxburghii, C. siamea, Cercis siliquastrum (Judas tree), Delonix
alata (white Gulmohar), D. regia (Gulmohar or Royal Poinciana), Parkinsonia aculeata
(Vilayati Kikar) and Saraca indica (Ashok tree).
• Medicinal Value: (1) Caesalpinia bunduc seeds are used in diarrhoea and rheumatism, (ii)
Cassia acutifolia leaves are used as purgative, (iii) C. alata leaves are used in skin infec-
tions, (iv) C. fistula fruit pulp is taken as a purgative. (v) C. occidentalis seeds and leaves
are used in skin infections; (vi) C. sophera leaf decoction is beneficial in acute bronchitis.
• Haematoxylin: The best known nuclear stain, haematoxylin, is obtained from the heart-
wood of Haematoxylon campechianum (logwood).
• Timber: Valuable timber, useful in making agricultural implements, is obtained from
Cassia fistula, Copaifera pubiflora, Hardwickia binata (Anjan), Hymenaea courbaril and
Tamarindus indica (Tamarind).
• Tannin: Tannin of commercial importance is obtained from the bark and seeds of Bauhinia
malabarica, B. purpurea, B. racemosa, B. vahlii, B. variegata, Caesalpinia coriaria, C.
digyna and C. sepiaria.
• Edible Products: (i) Floral buds of Bauhinia purpurea (Kachnar) are cooked as vegetable,
(ii) leaves and tender seeds of B. malabarica are also eaten, (iii) roasted seeds of Cassia
anther style
ovary lobe
stamen
petal stigma
filament
staminode sepal
pedicel A Stamen
An Open Flower ovary
stamen ovary locule

ovary wall
petal
pedicel
ovule
Gynoecium
sepal T.S. Ovary

pedicel
stem
L.S. Flower
leaflet
leaf
inflorescence
axis
flower
floral
buds
Fig. 21.49 Cassia fistula L.
occidentalis are used in place of coffee while that of Gymnocladus canadensis are used
as Kentucky coffee, (iv) acidic fruit pulp of Tamarindus indica is used for sauces, curries,
chutneys, etc.
• Gum is obtained from Bauhinia racemosa, B. variegata and Copaifera pubiflora.
• Fibre for making ropes is obtained from the bark of Bauhinia racemosa and B.
tomentosa.
• Red dye for making red ink is obtained from the heartwood of Caesalpinia sepiaria.
306 Plant Taxonomy
21.36.7 Description of a Common Representative

Cassia fistula Linn. (Amaltas; Fig. 21.49)
Habit: An ornamental cultivated tree. Leaf: Stipulate, petiolate, unipinnate and paripinnately com-
pound; leaflets 6–9, opposite, ovate, entire, acute, unicostate reticulate. Inflorescence: Racemose
raceme. Flower: Bracteate, pedicellate, zygomorphic, hermaphrodite, pentamerous, hypogynous.
Calyx: 5 sepals, free, petalloid (yellowish-green), quincuncial or imbricate, odd sepal anterior. Corolla:
5 petals, polypetalous, ascending imbricate, yellow, posterior petal small. Androecium: 10 stamens,
polyandrous; 3 anterior stamens large, 4 stamens of inner whorl medium-sized, and 3 posterior sta-
mens reduced into staminodes; anthers dithecous, dorsifixed, introrse. Gynoecium: MonocarpelIary,
superior, unilocular, marginal placentation; ovary sickle shaped; stigma reduced and curved. Fruit:
Legume.
Floral Formula: Br, , , K5, C5, A3+4+3 staminodes, G1.
21.37 SUBFAMILY PAPILIONOIDEAE PEA FAMILY OR BEAN FAMILY


Herbs, shrubs, or trees; leaves simple or imparipinnately compound or palmate; flowers zygomor-
phic; corolla papilionaceous (i.e. consisting of a standard, 2 wings and a keel); corolla aestivation
descending-imbricate; posterior petal outermost; stamens monadelphous or diadelphous.

Arachis, Cicer, Crotalaria, Dalbergia, Dolichos, Glycine, Indigofera, Lathyrus, Pisum, Vicia,
Vigna.

The subfamily Papilionoideae contains about 500 genera and over 10,000 species. It is distributed
all over the globe excepting arctic regions, and is largely represented in warm temperate regions of
both the Northern and Southern Hemispheres. Over 100 genera and 800 of its species have so far
been reported from India. Astragalus (Milk Vetch), the largest genus of this subfamily, contains
about 2,000 species. Some of the other larger genera, of which 100 or more species have been
reported, include Indigofera (700, Indigo), Crotalaria (600, Rattlebox), Desmodium (450, Beggar-
lice), Dalbergia (300, Shisham), Trifolium (300, Clover), Phaseolus (240, Cultivated beans), Lupinus
(200, Lupine), Vicia (150, Broad bean), Lathyrus (130, Sweet pea), Psoralea (130, Snakeroot),
Medicago (110, Alfalfa), Erythrina (100, Coral bean), and Vigna (100, Southern pea).
Takhtajan (1969) treated subfamily Papilionoideae as an independent family of order Fabales but
instead of Papilionaceae he calls the family as Fabaceae. Cronquist (1981) and Jones and Luchsinger
(1987) also used the name Fabaceae for Papilionaceae.

General Habit Mostly herbs; however, shrubs, trees, and climbers are also common; Clitoria,
Dolichos, Phaseolus, etc. are twiners; Crotalaria juncea is an example of shrub; Millettia is a
woody climber; Sesbania is a small tree whereas species of Dalbergia, Erythrina, Pterocarpus, etc.
are large trees.
Roots Lateral branches of roots of most Papilionoideae contain nodules or tubercles, which contain
nitrogen-fixing bacteria (Rhizobium) having the ability to fix the atmospheric nitrogen.
Leaves Usually alternate, stipulate, compound, with pulvinous leaf base; stipules are adnate
(Medicago, Trifolium), or free lateral (Cajanus, Sesbania), or foliaceous (Pisum); compound leaves
may be unifoliate (Desmodium gangeticum), or bifoliate (Zornia), or trifoliate (Medicago, Trifolium,
Trigonella); paripinnate (Sesbania), or imparipinnate (Robinia); in Indigofera cordifolia and I. lini-
folia, the leaves are simple; terminal leaflet modifies into a tendril in Lathyrus, Pisum and Vicia.
Inflorescence Usually racemose raceme (Melilotus alba), or head or capitulum (Trifolium), or spike
(Uraria), rarely an umbel, or panicles (Dalbergia); sometimes the flowers are solitary (some species
of Lathyrus).
Flowers Bracteate, sometimes bracteolate (Sesbania, Astragalus), complete, hermaphrodite, zygo-
morphic, pentamerous; hypogynous or sometimes perigynous.
Calyx 5 sepals, more or less united in a tube and persistent; valvate or imbricate; invariably the
odd sepal is anterior in position.
Corolla 5 petals, very unequal and papilionaceous (i.e. 1-posterior petal is largest and called stan-
dard or vexillum, 2 lateral petals are lanceolate and slightly curved and called wings, and 2 anterior
petals are asymmetrical and more or less united to form a boat-shaped structure and called keel);
aestivation is vexillary or descending-imbricate i.e. standard covers the wings, and the wings cover
the keel; all petals have a claw at the base.
Androecium 10 stamens, diadelphous, usually 9 + 1 (Fig. 21.50) or sometimes 5 + 5; in case of 9 + 1
arrangement, 9 stamens are united and 1 is free, and the free stamen is always posterior in position;
often the posterior stamen is sterile or even absent (Arachis hypogaea, Dalbergia sissoo) and the
remaining 9 stamens show monadelphous condition; anthers are dithecous, introrse and dehisce by
longitudinal slits; rarely all the 10 stamens are free (Sophora).
Gynoecium Almost same as in Mimosoideae and/or Caesalpinioideae.
Fruit and Seed Fruit usually a legume, sometimes indehiscent; rarely a lomentum (Desmodium);
matures inside the ground in Arachis hypogaea. Seeds usually with food reserves in cotyledons.
General Floral Formula Br, , , K(5), C1+2+(2), A(9)+1, G1.
308 Plant Taxonomy
vexillum or
standard calyx
seed
petal wing or
alae
keel
sepal
Various Corolla Lobes
Fruit
pedicel
A Flower
(viewed from anterior side) stigma stamen (1) stigma
stamens
(9)
ovary style
Stamens and Pistil
stamen
ovary petal
pedicel sepal
L.S. Flower
flower
tendril
leat
stem
Fig. 21.50 Lathyrus odoratus L.

Subfamily Papilionoideae is of great economic importance as a source of high-protein food
(pulses), ornamentals, oil, forage, timber, dye, etc. Some of the universally known economically
important plants of this subfamily include Glycine max (soybean, source of oil and high-protein
meal), Pisum sativum (garden pea), Cicer arietinum (gram or Chana), Medicago sativa (alfalfa,
world’s best forage crop), Arachis hypogaea (groundnut, source of edible seeds, oil, peanut butter),
Indigofera tinctoria (Indigo plant, source of indigo dye), and Lathyrus odoratus (sweet pea, well-
known ornamental).
• Pulses and Vegetables1—Source of Proteins and Starch of our Food: These include (i) Arhar
or pigeon pea (Cajanus cajan), (ii) Bankla or field bean (Vicia faba), (iii) Chana or gram
(Cicer arietinum), (iv) Gwar or cluster bean (Cyamopsis tetragonoloba), (v) Lobia or lima
bean (Phaseolus lunatus), (vi) Masoor or lentil (Lens culinaris syn. L. esculenta), (vii) Matar
or pea (Pisum sativum), (viii) Moth or dew gram (Vigna aconitifolia syn. Phaseolus aco-
nitifolius), (ix) Mung or green gram (Vigna radiata syn. Phaseolus radiatus), (x) Rajmah or
cowpea (Vigna unguiculata), (xi) Sem or bean (Lablab purpureus syn. Dolichos lablab), (xii)
Soybean or Soya (Glycine max syn. Glycine soja), (xiii) Urd or black gram (Vigna mungo
syn. Phaseolus mungo), and (xiv) Vilayati Sem or kidney bean (Phaseolus vulgaris).
• Ornamental Plants2 —Source of Beautification: Clianthus dampieri (Glory pea), Clitoria
ternatea (Butterfly pea, blue-flowered climber), Erythrina indica (Indian coral tree, orange
red-flowered), Lathyrus odoratus (sweet pea, variously-coloured flowers), Lupinus hirsutus
(Lupine, variegated-flowered annual), Robinia pseudoacacia (Black locust, white-flowered
tree), Sesbania grandiflora (Sesban), S. sesban (Jait), Sophora japonica (Japanese Pagoda
tree, multicoloured flowers), and Wisteria sinensis (Chinese Wisteria, variously-coloured
flowers).
• Oils: Arachis hypogaea (groundnut or peanut or Moongphali; source of edible peanut oil;
seeds are eaten after roasting; oil is used for preparing soaps, cosmetics; oil cake is used as a
fodder), Pongamia pinnata (Pongam oil; oil obtained from its seeds is used for illumination,
soap preparation and medicinal purposes), Psophocarpus tetragonolobus (Goa bean; seeds
yield the oil used for soap making and cooking), and Spatholobus roxburghii (oil from its
seeds is used for anointing and cooking).
• Fodder: Several plants of this family are of high forage value and grown as fodder for
cattles. These include Medicago sativa (Alfalfa or Rizka), M. abscura, M. hispida, Trifolium
repens (Clovers), T. pratense, Dolichos biflorus, Phaseolus aconitifolius, Pisum sativum,
Vicia benghalensis and V. sativa.
• Timber: Valuable timber used for furniture, cabinet works and other building materials is
obtained from Baphia nitida, Dalbergia sissoo (Shisham), D. latifolium (Kala Shisham), D.
melanoxylon (African black wood), Pterocarpus indicus (Malay Padauk), P. dalbergioides
(Andaman redwood), P. marsupium (Indian Kino tree), and P. santalinus (Red sandalwood,
or Lal Chandan).
• Medicinal Value: Several plants3 of this subfamily are used as the source of drugs. Some of
them, along with their parts utilized and the name of the disease, to be cured, in parenthesis,
include (i) Abrus precatorius (Ratti; leaves and roots in cough and cold); (ii) Crotalaria albida
(roots used as purgative); (iii) Glycyrrhiza glabra (Mulhatti or liquorice; roots in cough and
sore throat); (iv) Krameria triandra (roots in chronic diarrhoea); (v) Moghania grahamiana
1
Arranged alphabetically in order of their Hindi local names.
2
Arranged alphabetically in order of their botanical names.
3
Botanical names of plants arranged alphabetically.
310 Plant Taxonomy
(roots are anthelmintic); (vi) M. strobilifera (roots induce sleep); (vii) Physostigma veneno-
sum (seeds in eye troubles); (viii) Psoralea corylifolia (seeds used in leucoderma and lep-
rosy), (ix) Teramnus labialis (entire plant used in tuberculosis); and (x) Uraria lagopodioides
(leaves and roots used in intermittent fever).
• Insecticides: Roots of Derris elliptica, D. ferruginea and D. trifoliata, roots and seeds
of Milletia extensa, entire plant of Spatholobus roxburghii, and roots of Tephrosia vogelii
possess insecticidal properties.
• Sola Hats and Toys: Stuffing material of sunhats is the soft wood of Aeschynomene aspera
and A. indica. Pith of these plants is used for making toys.
• Dyes1: (i) Orange-red dye is obtained from the flowers of Butea monosperma (Dhak);
(ii) blue dye is obtained from the flowers and seeds of Clitoria ternatea; (iii) black dye is
produced from the flowers of Crotalaria striata; (iv) red dye is obtained from the flowers
of Erythrina variegata; (v) indigo, the famous blue dye, used for dyeing cotton clothes and
in making paints and printing ink, is obtained from Indigofera tinctoria and some other
species; (vi) yellow dye is obtained from the pods of Psoralea pricata; and (vii) red dye is
produced from the wood of Pterocarpus santalinus.
• Green Manuring: Because of the presence of nitrogen-fixing bacteria in the lateral roots,
several members of this subfamily are grown for enriching the soil with nitrogen and then
ploughed as “green manure” in the field. Some of such members include Crotalaria juncea,
Medicago sativa, Tephrosia vogelli, Trifolium pratense and Trigonella foenum-graecum.
• Fibre: Some of the fibre-yielding plants2 of this subfamily, used for making cords, ropes,
bags, etc. include Butea monosperma (Dhak), Crotalaria juncea (Sunnhemp or Sani),
Erythrina variegata, Millettia auriculata, Sesbania bispinosa (Dhaincha), S. cannabina and
S. sesban.
• Gums and Resins: Gums and resins obtained from some of the plants of this subfam-
ily along with their utility in parenthesis include Astragalus gummifer (in confectionery,
cosmetics and textile industry), A. strobiliferus (in glazing and calico-printing), Cyamopsis
tetragonoloba (in food and textile industry), and Myroxylum balsamum (in medicine and
perfumery).
• Honey Production: Lathyrus odoratus, Medicago sativa, Pisum sativum and Trifolium
pratense are some of the plants of this subfamily which are used for artificial culture of
bees for honey production.

1. Lathyrus odoratus Linn. (Sweet pea or Phool Matar, Fig. 21.50)
Habit: Annual, cultivated, garden ornamental, climbing herb. Leaf: Alternate, petiolate; stipulate,
stipule leafy; pinnately compound, imparipinnate; upper leaflets modify into tendrils. Inflorescence:
Racemose or solitary axillary flowers. Flower: Bracteate, complete, hermaphrodite, zygomorphic,
pentamerous, perigynous. Calyx: 5 sepals, fused, odd sepal anterior. Corolla: 5 petals, papilionaceous
1
Botanical names of plants arranged alphabetically.
2
Botanical names arranged alphabetically.
corolla (consisting of 1 standard or vexillum, 2 wings or alae and 2 fused petals forming keel or
carina); descending imbricate. Androecium: 10 stamens, diadelphous with 9 stamens united and
10th posterior stamen free, dithecous, basifixed, introrse. Gynoecium: Monocarpellary, semi-inferior,
unilocular, marginal placentation. Fruit: Legume.
Floral Formula: Br, , , K(5), C1+2+(2), A(9)+1, G1–.
2. Crotalaria medicaginea Lamk. (Fig. 21.51)
Habit: Annual herb. Leaf: Alternate, petiolate; stipulate, stipules free-lateral; trifoliate; leaf base
pulvinus; obovate, apex emarginate. Inflorescence: Racemose raceme. Flower: Same as in Lathyrus.
Calyx: Same as in Lathyrus. Corolla: Same as in Lathyrus. Androecium: 10 stamens, monadelphous,
dithecous, dorsifixed, introrse. Gynoecium: Almost same as in Lathyrus. Fruit: Legume.
Floral Formula: Br, , , K(5), C1+2+(2), A(10), G1–.
stigma stigma
style
style
stamen
keel ovary
pedicel wing
ovary
ovules pedicel
L.S. Flower
anther lobe Gynoecium

connective
flower
filament
Stamen
leaflet
petiole
leaf
stem

Fig. 21.51 Crotalaria medicaginea Lamk.
312 Plant Taxonomy
21.38 ROSACEAE ROSE FAMILY


Perennial herbs, shrubs, or trees; leaves stipulate; flowers pentamerous, actinomorphic; stamens
numerous and usually in multiple of 5; ovary superior to inferior; fruit various types; seeds non-
endospermic.

Rosa, Pyrus, Prunus, Eriobotrya, Potentilla, Rubus, Spiraea.

A family of approximately 100 genera and 3000 species which are cosmopolitan in distribution
but particularly abundant in Eastern Asia, North America and Europe. About 25 genera and 230
species of Rosaceae have so far been reported from India. Some of the largely represented genera
along with their common names and approximately reported species include Potentilla (Cinquefoil,
500), Prunus (peach, plum, almond, cherry, apricot, 430), Rosa (Rose, 250), Rubus (Raspberry,
blackberry, 250), Alchemilla (250), Crataegus (Hawthorn, 200), Sorbus (Chokeberry, 100), Acaena
(100), and Spiraea (100).
Named after the universally famous flowering plant, rose, the Rosaceae show great diversity in
types of fruit (Fig. 21.52).

General Habit Mostly trees (Prunus, Pyrus, Eriobotrya) and shrubs (Spiraea), and only some are
herbs (Fragaria indica, Potentilla supina); some are prickly climbers (Rosa moschata, Rubus) or
prickly shrubs (Rosa damascena).
Stem Herbaceous or woody, usually with thorns or prickles.
Leaves Mostly alternate, simple (Prunus, Nuttalia) or compound; usually stipulate, stipules paired
and sometimes adnate to the petiole (Rosa, Rubus), sometimes exstipulate (Spiraea); compound
leaves may be palmately compound (Fragaria) or pinnately compound (Rosa); extrafloral nectaries
present on the back of leaves in Prunus laurocerasus.
lnflorescence Highly variable from solitary flowers to racemose and cymose clusters; solitary (Rosa),
racemose raceme (Agrimonia, Prunus mystifolia), corymb (Spiraea), head or spike (Poterium), cymes
(Spiraea, Fragaria), or thyrsoid panicles (Eriobotrya).
Flower Mostly bracteate, bracteolate; usually actinomorphic but zygomorphic in Hirtella and
Parastemon, usually bisexual but unisexual in Poterium sanguisorba and Pygeum, usually complete
achenes
Filipendula
Rubus
Dryas
Alchemilla
Geum
Fragaria
Agrimonia
Potentilla
Mespilus
Fig. 21.52 Types of fruit in Rosaceae.
but apetalous in Alchemilla, pentamerous; usually perigynous (Rosa) or epigynous (Pyrus) and only
seldom hypogynous (Prunus); epicalyx usually present; usually the floral receptacle along with the
basal adnate portions of perianth is variously modified into hypanthium; the hypanthium is often
lined with a nectariferous disc.
314 Plant Taxonomy
Epicalyx Often present in several genera (Fragaria, Geum, Potentilla supina) and alternate with
calyx lobes.
Calyx 5 sepals, free or basally connate; valvate or imbricate; sepals often foliaceous (Rosa).
Corolla 5 petals, usually attached to the rim of hypanthium; petals are numerous, large and vari-
ously coloured in cultivated species of Rosa; petals absent in Alchemilla and Poterium.
Androecium Usually numerous, distinct stamens in whorls of 5, sometimes only 5 or 10 stamens;
anthers basifixed or dorsifixed, dithecous, introrse and dehisce by longitudinal slits; usually a honey-
secreting disc is present in between stamens and carpels. Undermentioned are some of the commonly
seen arrangements of stamens in some Rosaceous genera, though there may also be some deviations
from these arrangements:
1. Only 1 stamen in Alchemilla arvensis (Fig. 21.53A).
2. Only 4 stamens alternating with sepals in Alchemilla vulgaris (Fig. 21.53B).
3. Only 10 stamens with their filaments joined together (monadelphous) in Acioa guianensis.
4. Only 15 stamens arranged in an outer whorl of 5 pairs of antisepalous stamens and an inner
whorl of 5 antipetalous stamens in Nuttalia.
5. Only 20 stamens arranged in an outer whorl of 5 antisepalous pairs followed by a middle whorl
of 5 antipetalous and an inner whorl of 5 antisepalous stamens in Pyrus (Fig. 21. 53C).
6. Only 25 stamens arranged in an outer whorl of 5 antipetalous pairs, a middle whorl of 5 anti-
sepalous pairs and an inner whorl of 5 antipetalous stamens in Potentilla (Fig. 21.53D).
7. Only 30 stamens arranged in an outer whorl of 5 antisepalous pairs followed by a middle
whorl of 10 antipetalous and an inner whorl of 10 antisepalous stamens in Prunus.
8. Only 30 stamens arranged in an outer whorl of 5 antipetalous pairs followed by a 2nd and
3rd whorl of 10 stamens each in antisepalous and antipetalous positions in Spiraea.
9. Numerous stamens arranged in an outer whorl of 5 antipetalous pairs followed by numerous
whorls of 10 stamens each in alternating positions in Rosa (Fig. 21.54).
Gynoecium Highly variable from 1 to numerous pistils; apocarpous to syncarpous; present at the
base of the floral cup, or on the sides of floral cup, or enclosed by the floral cup; pistils are either
free from the floral cup or adnate to the floral cup; ovary superior to inferior with intermediate
stages; styles and stigmas as many as carpel number.
Monocarpellary (Alchemilla, Chrysobalanus, Prunus), or pentacarpellary and apocarpous
(Spiraea), or multicarpellary and apocarpous (Fragaria, Rosa, Rubus), or 5 carpels fused only in
the basal region (Nuttalia), or carpels united completely with each other as well as with the floral cup
(Pyrus); if the receptacle is convex or dome-shaped, the ovary is superior (e.g. Fragaria, Potentilla,
Rubus); if receptacle is cup-shaped and the carpels are developing on its inner surface, the ovary
is half-inferior (e.g. Prunus, Rosa, Spiraea); and if the carpels are fused with one another and also
with the floral cup (e.g. Pyrus), the ovary is inferior; placentation is marginal or axile.
A
B
C D
Fig. 21.53 A–D Floral diagrams of Alchemilla arvensis (A); A. vulgaris (B); Pyrus (C) and
Potentilla (D).
Fruit Various types (Fig. 21.52), varying from drupe (Prunus), pome (Pyrus, Malus, Crataegus,
Sorbus), follicles (Spiraea, Sorbaria, Physocarpus), etaerio of achenes (Potentilla, Fragaria, Rosa),
or etaerio of drupes (Rubus).
Seed Endosperm usually absent.
General Floral Formula Br, ≈, , K5 or (5), C5, Aμ, G1 – μ –, or G1 – μ, or G(μ)–.

• Edible Fruits: Variety of delicious fruits belong to Rosaceae, and this family should better
be called as the “family of edible fruits”. Some of them1 include almond or Badam (Pyrus
1
Arranged alphabetically according to their English names along with the Hindi names of some of them.
316 Plant Taxonomy
stamen
petal
carpels
sepal
pedicel
L.S. Flower
Floral Diagram
achenes
calyx tube
V.S. Fruit
floral bud
flower
leaflets
leaf
stem stipule
Flowering Branch
Fig. 21.54 Rosa indica L.
amygdalus), alpine strawberry (Fragaria vesca), apple or Seb (Pyrus malus), apricot or
Khubani or Zardalu (Prunus armeniaca), blackberry or Vilaiti Alucha (Rubus fruticosus),
blackcherry (Rubus molucannus), Ceylon raspberry (Rubus laciocarpus), cherry laurel
(Prunus laurocerasus), Chinese pear (Pyrus pyrifolia), dewberry (Rubus caesius), Japanese
plum (Prunus saticina), Loquat (Eriobotrya japonica), peach or Aadu (Prunus persica),
pear or Nakh (Pyrus communis), plum or Aloocha or Alubukhara (Prunus domestica sub.
sp. instita), quince (Cydonia oblonga), raspberry (Rubus idaeus), rowan (Sorbus aucuparia),
sour cherry or Alu-balu (Prunus cerasus), strawberry (Fragaria chiloensis and F. vesca),
sweet cherry (Prunus avium), wild Himalayan cherry or Padam (Prunus cerasoides) and
yellow Himalayan raspberry (Rubus ellipticus).
• Ornamental Plants: Rose (Rosa), the best ornamental plant of the world and represented
by its innumerable varieties, belongs to this family. Some of the commonly grown species of
Rosa include R. alba, R. banksiae, R. centifolia, R. chinensis, R. damascena, R. indica and
R. moschata. Some other garden ornamentals of Rosaceae are Acioa guianensis, Crataegus
mollis, Exochorda racemosa and several species of Physocarpus, Potenti11a, Sorbus and
Spiraea.
• Medicines and Perfume: (i) ‘Gulkand’, prepared from rose (Rosa) petals is a good tonic,
laxative as well as useful in tonsilitis; (ii) root-stocks of Geum urbanum are used in diar-
rhoea and chronic dysentery; (iii) female flowers of Hagenia abyssinica are the source of
an antihelmintic drug, cusso; (iv) decoction of leaves of Potentilla anserina is used against
arthritis and kidney stones; (v) dried bark of Pyrus pyrifolia is useful in cough and cold; (vi)
valuable perfume is obtained from the petals of Rosa damascena and other rose species.
• Bark and Wood: Bark of soaptree (Quillaja saponaria) is powdered and used in sham-
poos and hair tonics, while walking sticks and tool handles are prepared from the wood of
Docynia indica, Prunus cerasoides, Pyrus pashia and P. communis.

Rosaceae, included under order Rosales of series Calyciflorae of subclass Polypetalae by Bentham
and Hooker, are perhaps allied to Calycanthaceae in the order Laurales and to Myrtaceae (Hickey
and King, 1988). Some believe Rosaceae to be allied to Saxifragaceae. Hutchinson believed that
Rosaceae is derived from the same stock as Dilleniaceae. Rosaceae is also related to Leguminosae, the
flowers of Chrysobalanoideae and Prunoideae having monocarpellary pistil and in Chrysobalanoideae
the flowers being zygomorphic.
Fockey in Pflanzenfamilien divided Rosaceae into 6 subfamilies (Spiraeoideae, Pomoideae,
Rosoideae, Neuradoideae, Prunoideae and Chrysobalanoideae) whereas Hutchinson divided Rosaceae
into 20 tribes. However, majority of the recent workers (Jones and Luchsinger, 1987; Hickey and
King, 1988, etc.) divide Rosaceae into 4 well-marked subfamilies as under:
1. Spiraeoideae (fruit a follicle or capsule; carpels usually 2–5, free or connate at base, not
sunk in hypanthodium), e.g. Spiraea, Sorbaria, Physocarpus.
2. Maloideae (fruit a fleshy pome; ovary inferior, carpels 2–5, united to inner wall of hypan-
thium), e.g. Pyrus, Malus, Crataegus, Sorbus.
3. Rosoideae (fruit often dry; gynoecium of usually 10 or more pistils), e.g. Rosa, Potentilla,
Fragaria, Rubus, Alchemilla.
4. Prunoideae (fruit drupe; carpel usually 1, free of hypanthium), e.g. Prunus.
318 Plant Taxonomy
21.39 SAXIFRAGACEAE SAXIFRAGE FAMILY


Mostly perennial herbs; leaves alternate, usually deciduous and exstipulate; flowers pentamerous;
ovary superior to inferior, often half-inferior; fruit usually a capsule or berry; seeds with abundant
endosperm.

Ribes, Hydrangea, Saxifraga, Deutzia.

A family of about 80 genera and 1250 species (Hickey and King, 1988), Saxifragaceae are almost
cosmopolitan in their distribution. They are, however, found mainly in north-temperate and arctic
regions. Some of the larger genera are Saxifraga (370; Fig. 21.55), Ribes (150), Hydrangea (80),
Philadelphus (75), Escallobonia (60), Chrysoplenium (55), Parnassia (50) and Deutzia (50).

General Habit Perennial herbs and shrubs, and a few small trees; only rarely annual herbs; in
Saxifraga urbium the stem is short, procumbent and rooting at the nodes.
Leaves Simple, alternate and exstipulate; compound (Astilbe), deeply lobed (Francoa); opposite
(Hydrangea); usually deciduous; fleshy, spathulate to obovate to form basal rosettes in Saxifraga
urbium.
Inflorescence Usually racemose or cymose; rarely solitary; panicle with loosely arranged flowers
in Saxifraga urbium.
Flower Usually bisexual and actinomorphic; sometimes unisexual (Ribes) and zygomorphic (some
species of Saxifraga); receptacle flat or hollowed to various depths so that perianth lobes and stamens
may be perigynous or epigynous.
Calyx Usually 5 sepals, sometimes 4; poly- or gamosepalous; sometimes deeply coloured and
petaloid.
Corolla Petals usually of same number as that of sepals, polypetalous; developing on a receptacle
or on an hypanthium; alternisepalous; sometimes absent.
Androecium Stamens usually of same number as that of petals, or twice as many to the petals;
alternipetalous or obdiplostemonous; free; anthers usually dithecous, dehiscing longitudinally; nectar
glands or staminodes often present (Parnassia).
Gynoecium Bicarpellary, syncarpous; uni- to pentalocular, ovary superior to inferior, often half-
inferior; axile placentation, but sometimes parietal; numerous anatropous ovules; styles and stigmas
as many as the carpels.
stamens
(obdiplostemonous)
petals
flowers ovary
persistent
filaments
glandular
pedicel hairs
fruit C
A Bisexual Flower
persistent
calyx stigmas
H styles
A Fruit
ovary
anther
leaves lobes
sepals
pedicel
filament E
stem Ovary (With Petals and

Stamens Removed)
stigma
D
roots A Stamen
style
A
petals ovules
A Flowering Plant
ovary
sepals
glandular
pedicel hairs F
G
pedicel
T.S. Ovary B L.S. Ovary
A Floral Bud
Fig. 21.55 Saxifraga urbium
Fruit A capsule or berry.

Seed With abundant endosperm surrounding a small embryo.
General Floral Formula Br, ≈, , K5, C5, A5 or 5 + 5, G (2).
320 Plant Taxonomy

Except for rock gardens (Saxifraga) or perennial ornamentals (Astilbe), Saxifragaceae are not of
much importance. Many species of Astilbe, Bergenia, Heuchera and Saxifraga are grown widely
as garden ornamentals. Ornamental flowering shrubs of Saxifragaceae belong to several species
of Deutzia, Escallonia, Hydrangea, Philadelphus and Ribes. Bush edible fruits of gooseberries
(Ribes) and white currant (R. sativum), black currant (R. nigrum) and red currants (R. rubrum)
are also of some importance.

Saxifragaceae shows close affinities with Rosaceae, and it is difficult to separate the two. Saxifragaceae
members, however, have (i) more abundant endosperm, (ii) exstipulate leaves instead of stipulate
leaves of Rosaceae, and (iii) lesser numbers of pistils and stamens than Rosaceae. Saxifragaceae also
shows close relationship with Crassulaceae. However, Crassulaceae differs from it (i) in the floral
parts of a regular numerical plant, and (ii) pistils subtended by scale-like glands.
Hickey and King (1988) placed all herbaceous members of Saxifragaceae in four subfami-
lies, viz. (i) Astilboideae (e.g. Astilbe), (ii) Saxifragoideae (e.g. Saxifraga), (iii) Francooideae (e.g.
Francoa) and, (iv) Parnassioideae (e.g. Parnassia) and all woody members in three subfamilies,
viz. (i) Ribesoideae (e.g. Ribes), (ii) Hydrangeoideae (e.g. Hydrangea), and (iii) Escallonioideae (e.g.
Escallonia), Earlier, Engler (1930) treated Saxifragaceae as composed of 15 subfamilies, of which
most of them were treated as independent families by several workers, including Hutchinson.
21.40 MYRTALES
1. Leaves simple, exstipulate, generally glandular-punctate; usually opposite and only sometimes
alternate.
2. Flowers actinomorphic, usually bisexual.
3. Ovary 2 to 5 or more carpelled, syncarpous, usually inferior.
4. Usually axile or sometimes parietal placentation.
5. Different members show transition from perigyny to epigyny.
6. Flowers show distinct development of an hypanthium.
7. Internally, the stem generally contains internal phloem or intraxylary phloem.
Bentham and Hooker included Combretaceae, Myrtaceae, Lythraceae and 3 more families under
Myrtales. Engler and Diels subdivided order Myrtiflorae (= Myrtales) into 4 suborders and 23
families, of which some major ones are Lythraceae, Punicaceae, Combretaceae, Myrtaceae and
Onagraceae.
Only Combretaceae, Myrtaceae and Lythraceae are discussed in this text.
21.41 COMBRETACEAE COMBRETUM FAMILY

Polypetalae, Calyciflorae, Myrtales.

Trees or shrubs, often lianous; leaves simple, exstipulate; tubular receptacle along with adnate tubular
base of calyx form hypanthium; ovules 4–6 in a single locule, all ovules suspended from the locule
apex by slender funiculi.

Combretum, Lumnitzera, Quisqualis, Terminalia.

Combretaceae contains about 20 genera and 600 species distributed mainly in the tropical and
subtropical countries of the world. 8 genera and about 45 species of this family have been reported
from India, chiefly from Assam and West Bengal. Combretum and Terminalia are two largest genera
of the family. Lumnitzera littoria is found as a mangrove plant in Sundarban and other tidal forests
of India.

General Habit A family of exclusively arborescent taxa, consisting of tall trees (Terminalia), or
woody twiners of lianas, such as Combretum and Quisqualis.
Stem Solid with large mucilaginous sacs and abundant tannin.
Leaves Simple, alternate or sometimes opposite (Quisqualis); exstipulate; margin entire.
Inflorescence Racemose, or paniculate, or simple or branched spikes (Quisqualis, Fig. 21.56); rarely
racemose heads (Anogeissus).
Flower Bracteate, bisexual (rarely unisexual); actinomorphic but occasionally zygomorphic; pen-
tamerous; epigynous; tubular floral receptacle along with the adnate tubular base of calyx form a
hypanthium; rarely flowers are tri- to octomerous.
Calyx Usually 5 sepals, fuse to form a tube; sometimes 4 or 8 sepals; valvate, rarely imbricate.
Corolla Usually 5 or as many petals as sepals, free; located alternately with sepals; valvate, imbri-
cate, or twisted (Quisqualis); petals absent in Terminalia (Fig. 21.57).
Androecium Usually 10 or twice as many as the petals, arranged in 2 alternate whorls; stamens of
the outer whorl are antisepalous; sometimes only 4 or 5 stamens; filaments inflexed in bud; anthers
dithecous, versatile, dehiscing longitudinally.
Gynoecium Carpels 4–5 or less, syncarpous; ovary inferior, unilocular; angular or ribbed, with as
many angles or ribs as calyx lobes; ovules 2–6, anatropous, pendulous from the top of the locule;
style slender and solitary; stigma pointed or capitate; ovary is surrounded by nectariferous disc.
322 Plant Taxonomy
stigma
petal
stamen
corolla tube
L.S. Ovaries
style
ovary
A Flower flower
L.S. Flower
A Stamen
leaf
Fruit
stem
Fig. 21.56 Quisqualis indica L.
Fruit and Seed Fruit leathery, one-seeded drupe; often winged. Seed non-endospermic; embryo with
folded or convolute cotyledons.
General Floral Formula Br, ≈, , K(5), C5, A5+5, G(4 – 5).

Family is of little importance for domestic purposes.
• Ornamental Value: Favourite ornamental plants of this family include Bucida buceras
(black-olive tree), few species of Combretum which are scandant shrubs, Quisqualis indica
(Rangoon creeper), and Terminalia arjuna (Arjun).
• Timber: Valuable timber used in construction work is

obtained from Anogeissus acuminata, A. latifolia, and
several species of Terminalia, such as T. alata, T. bialata,
T. catappa, T. ivorensis (indigo timber), T. myriocarpa, T.
paniculata, T. procera and T. tomentosa.
• Medicinal Value: (i) Bark of Terminalia arjuna is
highly valued as a cardiac tonic; (ii) Terminalia bellirica
(“Bahera” or Belleric myroblans) and T. chebula (“Hararh”
or black myroblans) are the two constituents of “Triphala”,
the famous Ayurvedic Indian preparation along with
Phyllanthus emblica (‘Aamla’) of family Euphorbiaceae as
its 3rd constituents; (iii) Leaves of Calycopteris floribunda
are used in dysentery and malarial fever; (iv) Leaves of
Combretum acuminatum are anthelmintic while that of C. Fig. 21.57 Floral diagram
roxburghii are used in malarial fever. of Terminalia bellirica.
• Tanning and Dyeing: The bark and fruits of Terminalia
alata, T. arjuna, T. catappa, T. chebula, T. ivorensis and T. citrina are used for tanning and
dyeing purposes.
• Edible Nut: Nuts of Terminalia catappa (Indian almond) are edible.

Bentham and Hooker placed Combretaceae under order Myrtales, series Calyciflorae and subclass
Polypetalae. Most of the other taxonomists (Takhtajan, 1969; Cronquist, 1981; Thorne, 1983) also
placed Combretaceae under order Myrtales. From the affinities point of view, Combretaceae is closely
related to Myrtaceae and Rhizophoraceae, and to some other families of Myrtales to some extent.
21.42 MYRTACEAE MYRTLE FAMILY

Large trees or shrubs; leaves glandular-punctate; stamens numerous with their anther connectives
gland-tipped; ovary inferior; axile or parietal placentation.

Callistemon, Eucalyptus, Myrtus, Psidium, Syzygium.

A family of about 140 genera and 3,400 species, distributed mainly in subtropical and tropical
regions of the world. Family is particularly abundant in Australia where some Eucalyptus trees are
324 Plant Taxonomy
world’s tallest angiosperms. Myrtaceae is represented in India by about 15 genera and over 170 spe-
cies. Beautiful pinkish flowers of Myrtus communis are considered highly sacred among Greeks,
Egyptians, Jews, and Persians, and used in religious rites and ceremonies. Some of the larger genera
of Myrtaceae along with their number of reported species include Eugenia (1000), Eucalyptus (500),
Syzygium (500), Myrcia (500), Psidium (140), Myrtus (100) and Melaleuca (100).

General Habit Trees and shrubs, varying from small creepers to giant Eucalyptus reaching to a
towering height of 300–400 feet; lysigenous cavities containing ethereal oils are present in young
stem, leaves, floral parts and fruits.
Leaves Usually opposite, but rarely alternate; simple; exstipulate; usually entire, gland-dotted;
evergreen; and coriaceous.
Inflorescence Usually cymose, sometimes racemose, rarely of solitary flowers; multichasial cyme
in Eucalyptus; spikes in Callistemon; solitary axillary in Psidium guajava and Myrtus communis;
racemose raceme in Barringtonia.
Flower Bracteate; often with 2 bracteoles; bisexual, actinomorphic, penta-or tetramerous; epigynous
(Eucalyptus, Psidium) or perigynous (Tristania); in several genera, the floral receptacle unites with
the gynoecium at the base and often prolonged into a hypanthium.
Calyx 4 or 5 sepals, free or united; valvate or quincuncial; sepals in Eucalyptus are thrown off
unopened as a lid; sepals contain oil glands.
Corolla 4 to 5 petals, free (Callistemon citrinus) or united (Eucalyptus); imbricate; often nearly
circular; in Eucalyptus citridora corolla fuses with calyx to form a cup called operculum (Fig. 21.58);
few Eugenia species lack petals.
Androecium Stamens numerous, free (Psidium), or sometimes basally connate (Callistemon); usu-
ally bent inwards in bud; connectives often gland-tipped; anthers usually 2-celled, versatile, introrse
and dehiscing longitudinally or sometimes apically; sometimes stamens are grouped into as many
clusters as petals and are obdiplostemonous (Syzygium); stamens variously coloured.
Gynoecium 2 to 5 or more carpels, syncarpous; ovary inferior or semi-inferior; with one to many
(often 2–5) locules; 2 to many anatropous or campylotropous ovules in each locule; usually axile
or rarely parietal placentation; style one, long and simple; stigma capitate.
A nectar-secreting, circular disc is present on the inside of the upper part of the floral cup.
Fruit and Seed Fruit variable; fleshy berry (Psidium, Eugenia), capsule (Callistemon, Eucalyptus),
nut (Darwinia), or drupe (Pimenta). Seeds with little or no endosperm.
Pollination and Dispersal Pollination entomophilous. Insects are attracted because of nectar and vari-
ously coloured stamens. Seeds are dispersed by wind, birds or other animals.
General Floral Formula Br, Brl, ≈, , K4–5 or (4–5), C4–5 or (4–5), Aμ, G (2 – 5).
stamens stigma
style
ovary
ovule operculum
pedicel
anther
L.S. Flower lobe
Floral Diagram
stigma
filament
Stamens style
pedicel
Gynoecium
inflorescence
flower buds
leaf
stem
Flowering Branch
Fig. 21.58 Eucalyptus citridora Hook.

• Edible Fruits: Psidium guajava (Amrood or Guava), P. cattleianum (Strawberry guava),
Eugenia uniflora (Surinam-cherry), Feijoa sellowiana (Feijoa), Myriciaria cauliflora
(Jaboticabo fruit) and Syzygium cumini syn. Eugenia jambolana (Jamun or Jambolan) are
some of the common delicious fruits. ‘Jamun’ fruits are useful for diabetic patients.
326 Plant Taxonomy
ovary wall stigma style
locule stamen
ovule
petal
placenta
ovary
ovule
T. S. Ovary sepal
L. S. Flower
flower
leaf stigma
fruit
style
Flowering Branch
ovary
Floral Diagram Gynoecium
Fig. 21.59 Callistemon lanceolatus D.C.
• Oil Plants: (i) Eucalyptus oil, obtained from Eucalyptus globulus, E. citridora and sev-
eral other species is used widely in medicine, perfumery and other industries, (ii) clove
oil is obtained from the dried flower buds of Syzygium aromaticum (Laung or clove tree);
(iii) cajuput oil is obtained from the leaves of Melaleuca leucadendron, (iv) oil of bay rum
is obtained from leaves and flower buds of Pimenta racemosa.
• Spices: Cloves (Laung) are the dried flower buds of Syzygium aromaticum while allspice
are the unripe berries of Pimenta dioica.
• Ornamental Plants: Hundreds of species of Eucalyptus are grown as ornamental plants.
Other garden ornamentals of Myrtaceae include Callistemon lanceolatum (bottle brush or
Laila Majnu plant), C. linearis, C. viminalis, Myrtus communis (Myrtle), Leptospermum
laevigatum (Australian tea tree), Melaleuca leucadendron (cajuput tree), and Tristania con-
ferta (Brisbane box).
• Timber: Valuable timber is obtained from several species of Eucalyptus, Eugenia,
Barringtonia and Coreya.
• Paper: Eucalyptus wood is used for paper manufacturing.

• Gum: Valuable gum is obtained from Eucalyptus globulus (blue gum), E. maculata (citron
gum) and E. rostrata (red gum).

Almost all taxonomists place Myrtaceae under order Myrtales. Myrtaceae is allied to Rhizophoraceae,
Lythraceae, Combretaceae and Onagraceae, and to some extent to Melastomaceae and
Sonneratiaceae.
Myrtaceae is divided into 2 subfamilies i.e. (i) Myrtoideae (fruit berry, rarely a drupe; leaves
always opposite), and (ii) Leptospermoideae (fruit dry, leaves opposite or alternate).

Callistemon citrinus Skeels syn. C. lanceolatus D.C. (Bottle brush or Laila Majnu, Fig. 21.59).
Habit: Large tree. Leaf: Simple, alternate, exstipulate, subsessile, lanceolate, gland-dotted.
Inflorescence: Long pendant spike. Flower: Bracteate, subsessile, complete, hermaphrodite, acti-
nomorphic, pentamerous, epigynous, crimson-yellow to red. Calyx: 5 sepals, fused, imbricate.
Corolla: 5 petals, polypetalous, imbricate or quincuncial. Androecium: Stamens numerous, filaments
long, bright red and united at the base to form a staminal sheath; dithecous, dorsifixed, introrse.
Gynoecium: Tricarpellary, syncarpous, inferior, trilocular, many ovules in each locule, axile placen-
tation. Fruit: Capsule.
Floral Formula: Br, ≈, , K(5), C5, Aμ (basally fused), G (3).
21.43 LYTHRACEAE (LOOSESTRIFE FAMILY)


Herbs to trees; leaves simple, opposite; hypanthium present; corolla crumpled; ovary superior; seeds
non-endospermic.

Lawsonia, Punica,1 Lagerstroemia and Woodfordia.

A family of about 25 genera and 550 species, Lythraceae are cosmopolitan in distribution. About
12 genera and over 50 species have been reported from India. Some of the larger genera along with
their approximately reported species are Cuphea (250), Lagerstroemia (43) and Lythrum (35).
1
Several taxonomists include Punica under a separate unigeneric family Punicaceae.
328 Plant Taxonomy

General Habit Herbs, shrubs (Lawsonia inermis, Woodfordia fruticosa), or trees (Sonneratia); rarely
spinescent (Lawsonia).
Leaves Usually opposite, or whorled; simple; exstipulate or with minute stipules; margin entire.
Inflorescence Racemose raceme, panicle (Lagerstroemia), or cyme (Punica).
Flower Bisexual, actinomorphic or zygomorphic (Cuphea); usually tetra- to hexamerous; hypan-
thium present; sepals appearing like lobes of hypanthium.
Epicalyx Hypanthium is sometimes subtended by the connate pairs of bracts representing epicalyx
(Lythrum).
Calyx 4 to 8 sepals; sometimes forming a hollow tube; valvate.
Corolla Petals of same number as sepals (i.e. 4 to 8), polypetalous, crumpled in bud (Fig. 21.60A);
petals insert on the margin of the receptacular cup and alternate with sepals; sometimes petals
absent (Rotala, Peplis).
Androecium Stamens typically twice as many as sepals (Fig. 21.60B), and only rarely fewer or
more; inserted often very low down on the calyx tube or hypanthium; stamens generally arranged
in 2 whorls; filaments unequal; anthers dithecous, introrse and dehisce longitudinally.
Gynoecium 2–6 carpels, syncarpous, 2–6 locular, numerous ovules in each locule, axile placenta-
tion; ovary superior or half-inferior; style simple, stigma capitate; some genera show heterostyly
(Lythrum).
Fruit and Seeds Fruit usually a dry capsule. Seeds non-endospermic.
General Floral Formula ≈, , K4–8 or (4–8), C4–8, A8–16, G (2–6) or (2–6)–.
A B
Fig. 21.60 Floral diagrams—A: Lythrum and B: Woodfordia fruticosa.

21.43.6 Economic lmportance

• Lawsonia inermis (Mehndi or Henna plant) leaves are the source of dye, henna, used
for staining fingers, nails, hairs, and also wool, silk and leather. Leaves and flowers of
Woodfordia fruticosa are also used for tanning and dyeing silk.
• Punica granatum (Pomegranate), a small tree, is cultivated for its edible juicy fruits.
• Timber for making boats and other similar articles is obtained from Lagerstroemia speciosa,
L. indica, L. lanceolata, etc.
• Ornamental plants of Lythraceae include Cuphea lanceolata (red-flowered), C. ignea (cigar-
flower), Punica granatum, Lawsonia inermis (hedge-plant), Lagerstroemia indica (pinkish
flowers), L. speciosa (purple flowers) and several species of Lythrum.

Lythraceae is closely allied to Myrtaceae, Onagraceae and Melastomaceae. Bentham and Hooker
treated genus Punica under Lythraceae. However, in several recent taxonomic treatments Punica is
placed in an independent unigeneric family Punicaceae.
21.44 PASSIFLORALES
1. Flowers regular; usually unisexual and only sometimes bisexual.
2. Flowers pentamerous, usually epigynous and only rarely perigynous.
3. Ovary is usually tricarpellary, syncarpous, inferior, unilocular or trilocular.
4. Placentation parietal or axile.
5. Fruit pepo or capsule and seeds non-endospermic.
Bentham and Hooker included Cucurbitaceae, Begoniaceae, Passifloraceae and four more fami-
lies under Passiflorales. Engler and Diels did not use the name Passiflorales, and treated only
Cucurbitaceae under order Cucurbitales, and Begoniaceae and Passifloraceae under order Parietales
along with 29 more families.
Only Cucurbitaceae, Begoniaceae and Passifloraceae have been treated in this text.
21.45 CUCURBITACEAE GOURD FAMILY

Polypetalae, Calyciflorae, Passiflorales.

Climbing or prostrate, tendril-bearing herbs; flowers usually unisexual and pentamerous; ovary
inferior; fruit pepo or berry.
330 Plant Taxonomy

Citrullus, Coccinia, Cucurbita, Lagenaria, Luffa, Momordica, Trichosanthes.

A family of about 110 genera and between 650 to 850 species, Cucurbitaceae are distributed mainly in
tropical and subtropical regions of the world. Its 37 genera and about 100 species have been reported
from India. Some of its largely represented genera along with their number of approximately reported
species include Momordica (45, bitter gourd), Cucumis (25, Cucumber, muskmelon), Cucurbita (15,
pumpkin, gourd, squash), Lagenaria (6, bottle gourd), Luffa (6, vegetable sponge or Loofah), and
Echinocystis (15, wild cucumber).

General Habit Annual or perennial prostrate herbs or vines usually with spirally coiled tendrils;
very rarely shruby (Acanthosicyos) or arborescent (Dendrosicyos).
Stem Herbaceous, usually pentangular with ridges and furrows; contains two rings of bicollateral
vascular bundles.
Leaves Alternate, exstipulate, simple but often palmately or pinnately lobed; long-petioled; palmately
veined; tendrils present (tendrils, interpreted variously as stipules, leaves, shoots, flower stalks, stems
or even as roots, are perhaps stipular in origin).
Inflorescence Axillary and basically cymose or the flowers solitary; plants are monoecious or
dioecious.
Flower Usually unisexual and only rarely bisexual (Schizopepon); actinomorphic; pentamerous;
epigynous or rarely perigynous (Actinostemma); female flowers contain staminodes while male
flowers contain pistillode.
(a) Male Flower
Calyx 5 sepals, tubular, gamosepalous, rarely distinct; fused to ovary wall in female flower; valvate
or imbricate.
Corolla 5 petals, usually united to form a campanulate or salver-shaped corolla; rarely petals are
free (Fevillea); usually petals alternate with the sepals; imbricate or valvate; yellow or white.
Androecium 1–5 stamens (typically 5, more or less united into a column, but very variable in struc-
ture or highly modified into a number of types of cohesion and twisting of anthers and filaments);
usually 5 stamens appear as 3 stamens, with 2 groups of 2 fused together forming 2 compound
stamens, plus 1 simple stamen. Some of the androecium types found in Cucurbitaceae are:
(i) In Fevillea, there are 5 free stamens alternating with polypetalous coralla.
(ii) In Thladiantha, filaments of 4 stamens coherent below in 2 pairs, the 5th stamen remaining
free.
(iii) In Sicydium, the condition is same as in Thladiantha but filaments of the pairs become
united at the base.
(iv) In Bryonia, Citrullus and Momordica, the coherent filaments and anthers of all the 5 sta-
mens unite completely so that the androecium appears to be made up of 3 stamens; of these,
2 have 4 cells due to complete union and the 3rd one has only 2 cells.
(v) In Cucurbita, the anthers are all united and spirally twisted, and filaments free below and
united above or completely united. Such folded anthers are called conduplicate.
(vi) In Cyclanthera the stamens completely unite with the lobes of the anthers forming two
horizontal rings surrounding a peltate mass of connectives.
Gynoecium Represented by rudimentary pistillodes.
(b) Female Flower
Calyx and Corolla Same as in male flower.
Androecium Represented only by staminodes, if any.
Gynoecium Usually tricarpellary, syncarpous, ovary inferior. According to Engler, Eichler, Wettstein
and Willis, the ovary is trilocular, the carpels intrude and meet at the centre and the placentation
is axile. However, Rendle opined that the ovary is unilocular and the placentation is parietal. In
some cases, carpels range from 1 to 10; ovules numerous and anatropous; style simple or 3-parted;
stigmas as many as carpels.
Fruit and Seed Fruit a pepo or capsule. Seeds often flat, numerous, with straight embryo and no
endosperm; seeds are winged in Zanonia.
General Floral Formulae
(a) Male Flower: ≈, , K(5), C (5), A5 or (2)+(2)+1, G0.
(b) Female Flower: ≈, , K(5), C (5), A0, G(3).

• Edible Fruits and Vegetables: Cucurbitaceae is a family of several edible fruits which are
either cooked or eaten raw in our daily food. Some of them nclude (i) Benincasa hispida
(petha-kaddoo or white gourd), (ii) Citrullus vulgaris (Tarbooj or watermelon), (iii) C.
vulgaris var. fistulosus (Tinda or round gourd), (iv) Coccinia cordifolia (Kanduri or Kovai
fruit), (v) Cucumis anguria (Gherkin), (vi) C. melo (Kharbooja or melon), (vii) C. melo
var. momordica (Phoont or Kachra), (viii) C. melo var. utilissimus (Kakdi). (ix) C. sativus
(Kheera or Cucumber), (x) Cucurbita maxima (Red pumpkin), (xi) C. moschata (Sitaphal or
pumpkin), (xii) C. pepo (Field pumpkin), (xiii) Lagenaria siceraria syn. L. vulgaris (Lauki
or bottle gourd), (xiv) Luffa acutangula (Kali Tori), (xv) L. cylindrica syn. L. aegyptiaca
(Ghia Tori), (xvi) Momordica charantia (Karela or bitter gourd), (xvii) M. cochinchinensis
(Kakrol or Bhat Karela), (xviii) Trichosanthes anguina (Chichenda or snake gourd), and
(xix) T. dioica (Palwal or Parwal).
• Ornamental Plants: Some species of Benincasa, Coccinia, Ecballium, Luffa, Sechium and
Trichosanthes are of ornamental value.
• Some Luffa species provide sponges for oil filters and bathing.
332 Plant Taxonomy
petal
tendril
stamen
sepal
pedicel
L.S. Male Flower
flower
petal
stigma leat
sepal
ovary A Stamen
ovary wall
pedicel stem
locule
Flowering Branch
L. S. Female Flower ovule
T.S. Ovary
Floral Diagram (female flower) Floral Diagram (male flower)
Fig. 21.61 Luffa cylindrica (L). Roem. syn. L aegyptiaca Mill.
• Musical instruments and some decorative articles are prepared from fruits of Lagenaria and
Benincasa.
• Seeds of Citrullus vulgaris and Cucumis melo are highly nutritious and used in preparing
sweets.

Systematic position of Cucurbitaceae is still highly controversial and its affinities are not very
clear among taxonomists. Its characters such as herbaceous nature, presence of bicollateral vascular
bundles, unisexual epigynous flowers, curved anther lobes, etc. make it a truly advanced family.
Engler, therefore, placed it in a monotypic order Cucurbitales and thought it to be closely allied to
Campanulaceae. Bentham and Hooker placed it under order Passiflorales. Rendle thought it to belong
to order Peponiferae while Takhtajan (1969) placed it under Cucurbitales in between Passiflorales and
Begoniales, and mentioned that Cucurbitaceae is “very closely related to the family Passifloraceae”.
Cronquist (1981) placed it under order Violales along with 23 more families while Thorne (1983)
placed it under suborder Cucurbitineae of order Violales along with 16 more families. Cronquist
considers Violales to be a natural taxon derived from Theales.

Luffa cylindrica (L.) M. Roem. syn. L. aegyptiaca Mill. (Ghia Tori; Fig. 21.61)
Habit: A tendril-bearing herb. Stem: Pentangular. Leaf: Alternate, simple, exstipulate, palmately
lobed. Inflorescence: Male flowers arranged in raceme clusters; female flowers solitary. Flower:
Bracteate, incomplete, actinomorphic, unisexual, pentamerous. MALE FLOWER—Calyx: 5 sepals,
fused, valvate. Corolla: 5 petals, gamopetalous or distinct, imbricate, yellow. Androecium: 5 stamens
appearing as if 3, of which 4 stamens are fused in 2 groups of 2 each, and 5th stamen is free.
Gynoecium: Absent or represented by rudimentary pistil. FEMALE FLOWER—Calyx and Corolla:
Same as in male flower. Androecium: Absent. Gynoecium: Tricarpellary, syncarpous, inferior, uni-
locular or divided incompletely into 3 locules, many ovules, parietal placentation; styles 3; stigma
forked. Fruit: Pepo. Floral Formulae: Same as mentioned in general description of family.
21.46 BEGONIACEAE BEGONIA FAMILY


Unisexual flowers; the staminate flowers are zygomorphic with numerous stamens arranged in many
whorls; the pistillate flowers contain inferior, angled or winged ovary with lobed or forked placentae,
generally twisted stigmas; seeds generally non-endospermic and contain oily embryo.
21.46.3 Common Indian Genus

Begonia.

Only 5 genera and about 920 species of this family have been reported so far. Begonia, with about 900
species, is largest genus distributed widely in tropics and subtropics. Hillebrandia found in Hawaii
334 Plant Taxonomy
and Symbegonia found in New Guinea are both monotypic genera. 3 species each of Begoniella and
Semibegoniella have also been reported.

General Habit Mostly perennial, monoecious herbs or low shrubs; erect, creeping or climbing with
succulent stems and thick rhizomes or tubers.
Stem Succulent, thick, rhizomatous or tuberous; somewhat jointed; in Begonia semperflorens the
stem is glabrous, fleshy, reddish-green (Fig. 20.62) attaining a height of 15 to 45 cm.
Leaves Simple, basal or alternate in 2 ranks; usually asymmetrical; stipulate, stipules large, membra-
nous and deciduous; petiolate, palmately-nerved; in B. semperflorens the leaves are roundish-ovate,
somewhat oblique at the base and have serrulate or ciliate margins.
Inflorescence Axillary dichasial cyme with unisexual flowers.
Flowers Unisexual; often zygomorphic, sometimes actinomorphic.
(a) Male Flower
Perinath 2, valvate, petalloid, free sepals, and 2 usually smaller, valvate petals.
Androecium Stamens 2 to numerous, in many whorls, free or basally connate; anthers dithecous,
basifixed.
Gynoecium Absent or rudimentary.
(b) Female Flower
Perinath 2 to many, imbricate, petalloid tepals not differentiated into colyx and corolla.
Androecium Absent.
Gynoecium Tricarpellary, syncarpous; inferior ovary; trilocular, with numerous anatropous ovules
on axile placentation; styles 2–5 free or basally connate; in Hillebrandia the ovary is half-inferior;
stigmas strongly papillose and often twisted.
Fruit Usually a loculicidal, winged capsule, rarely berry.
Seeds Numerous, minute, Non-endospermic with straight oily embryo.
Male flower , , P2+2, A2 – μ, G0.
Female flower ≈, , P2–μ, A0, G(3).

Except that of great horticultural or ornamental value of genus Begonia, the Begoniaceae contain
no plant of much economic importance. Large number of species of Begonia are grown as decora-
tive house plants. They are commonly grown for summer bedding in lawns of gardens.
staminate
flower
inferior
ovary
B
leaf
A Pistillate Flower
stem
A
A Flowering Shoot
(Upper Portion)
anthers
stigmas
(papillose)
styles
C G
Cluster of Stamens Basally Connate Styles
stigmas
(papillose)
connective
ovules
wing
filament locules
(3) ovules
D
A Single Stamen
wing ovary
E
T.S. Ovary
F
L.S. Inferior Ovary
Fig. 21.62 Begonia semperflorens Link and Otto.

Engler and Diels treated Begoniaceae under suborder Begoniineae of order Parietales, and Lawrence
(1951) also followed it. Bessey placed it under order Loasales along with families of two other
336 Plant Taxonomy
suborders (Datiscineae and Ancistrocladineae) of Parietales. Randle transferred Begoniaceae under

order Cucurbitales and treated it with the Cucurbitaceae. Bentham and Hooker treated Begoniaceae
under order Passiflorales of series Calyciflorae of Polypetalae and discussed it along with
Cucurbitaceae and five more families. Hickey and King (1988) described Begoniaceae under order
Violales of Dicotyledons.
21.47 PASSIFLORACEAE PASSION FLOWER FAMILY


Passifloraceae members can be recognised by the combination of the usually climbing habit, the
uniflowered peduncles often in pairs, the variously modified corona, androgynophores, and the seeds
with often mucilaginously pulpy aril.

Passiflora, Adenia.

A medium-sized family of 16 genera and over 650 species distributed mainly is tropical and warm
temperate regions. Of the three large genera, Passiflora (500) is distributed chiefly in USA, Asia
and Australia, while Adenia (92) is found mainly in tropical and South Africa, Indo-Malaysia and
South-West Arabia, and Typhostemma (35) in tropical and South Africa

General Habit Chiefly shrubs, or herbs; often climbers or lianous with axillary tendrils.
Stem Herbaceous, weak.
Leaves Alternate; simple or compound; stipulate; tendrillar, tendrils opposite the leaves; in Blue
Passion flower (Passiflora caerulea, Fig. 20.63) the leaves are palmate, 5- or 7-lobed with axillary
tendrils.
Inflorescence Modified dichasial cyme (Passiflora caerulea, Fig. 21.63), in which the central flower
is represented by a tendril, one of the lateral outgrowths develops into a flower, and the opposite
lateral outgrowth remains undeveloped.
Flowers Bracteate; bisexual or rarely unisexual; actinomorphic; axillary and usually in pairs.
Receptacle of various shapes, usually hollowed and generally with a central androgynophore; usu-
ally terminated by outgrowths of petalloid or staminodial appearance forming a fleshy, cup-shaped
corona. Puri (1948) interpreted that corona is made up of (i) the hypanthodium, and (ii) all associ-
ated structures situated between the perianth lobes and the stamens.
style
stigma
ovary
anther
filament
androgynophore
operculum pali and radii
of corona
tendril petal
sepal
C limen
hypanthium
L.S. Flower bracts
flower
leaf
tendril B
Flower (Anterior view)
stigma
A
style
A Flowering Branch stamen
locule ovary
ovules
E stigma
(capitate) Pali and radii of
T.S. Ovary corona
androgynophore petal
style sepal
persistent
bracts
F hypanthium
Style and Stigma bract
fruit
G D
A Fruit L.S. Central Part of Flower
Fig. 21.63 Passiflora caerulea L. (Blue Passion-flower).

338 Plant Taxonomy
Calyx 3–5 sepals, free or basally connate, often petalloid; imbricate; persistent.
Corolla 3–5 petals, free or basally connate, sometimes petals absent; imbricate.
Androecium 3–5 or more stamens, sometimes upto 10, usually opposite the petals, developing from
the base of corona or from receptacle rim or hypogynous from the apex of gynophore, thus forming
an androgynophore; anthers free or basally connate, dithecous, dehiscing longitudinally; some genera
show the presence of staminodes.
Gynoecium 3–5 carpels, syncarpous; ovary superior, unilocular, containing many anatropous ovules,
parietal placentation; pistil often raised on gynophore or more commonly on androgynophore; styles
as many as carpels, free or all connate; stigmas 3–5, often capitate or discoid.
Fruit A capsule or a berry.
Seeds With straight embryo, and fleshy aril and endosperm.
General Floral Formula Br, ≈, , K3–5 or (3 – 5), C3–5 or (3 – 5), A3 – 5 or (3 – 10), G(3 – 5).

• Ornamental plants of domestic importance include several species of Passiflora (e.g. P.
edulis, P. ligularis, P. laurifolia, P. caerulea, P. quadrangularis etc. Most of these species
also provide edible fruits. Several of these species are cultivated for their attractive flowers
and also grown in glasshouses.
• Passiflora edulis is widely known in tropical regions for its edible fruits (Passion fruit or
Purple Granadilla).
• Some species of Passifloraceae produce cyanide and are thus deadly poisonous. So, care
must be taken while eating the fruits of unidentified species.

Bentham and Hooker treated Passifloraceae under order Passiflorales of series Calyciflorae of
Polypetalae along with families like Cucurbitaceae and Begoniaceae. Hutchinson, however, treated
both Cucurbitaceae and Begoniaceae under order Cucurbitales while Passifloraceae under order
Passiflorales quite close to Cucurbitales.
21.48 FICOIDALES
1. Flowers usually solitary, showy, each with numerous undifferentiated floral parts.
2. Flowers mostly actinomorphic, bisexual and epigynous.
3. Numerous stamens arranged spirally or in clusters.
4. Ovary syncarpous, usually inferior, unilocular, parietal placentation.
5. Mostly prickly or spiny, fleshy succulent plants.

6. Seeds with scanty or no endosperm, often arillate with curved or straight embryo.
Ficoidales includes only Cactaceae and Ficoideae according to Bentham and Hooker. Engler placed
the single family Cactaceae in the Opuntiales, an order he considered to have been derived from the
Parietales. Ficoideae, also sometimes named as Aizoaceae, has been treated by Lawrence (1951) to
belong to order Centrospermae along with 9 more families, chief among which are Chenopodiaceae,
Amaranthaceae, Nyctaginaceae, Portulacaceae and Caryophyllaceae.
Only Cactaceae is discussed here. Families like Chenopodiaceae, Amaranthaceae and
Caryophyllaceae have been discussed elsewhere in this text.
21.49 CACTACEAE CACTUS FAMILY

Polypetalae, Calyciflorae, Ficoidales.

Mostly prickly or spiny, fleshy succulent plants; flowers usually solitary, showy, each with numerous
floral parts; ovary unilocular, inferior; parietal placentation.

Opuntia, Mammillaria, Cereus, Echinocactus.

Cactaceae includes approximately 85 genera and 2,000 species. But different authorities mention the
number of genera differently between 30 to 200. Members are distributed chiefly in semi-desert or
drier regions of tropical and subtropical America, and also in Africa, Mauritius, Seychelles, India,
Sri Lanka, and Australia. Mainly the Cactaceae are the succulents of New World. Main species
native of Old World is Rhipsalis. Some of the larger genera along with their number of reported
species are Mammillaria (200–300), Opuntia (250), Echinocereus (75), Rhipsalis (60), Cereus (50),
Cephalocereus (48), Echinopsis (35), Epiphyllum (21) and Pereskia (20).

General Habit Fleshy, succulent or xerophytic; perennial herbs, shrubs, or trees of diverse forms;
with spines or bristles or both.
Roots Root system generally shallow, with elongated, slender but fleshy roots.
Stem Fleshy, of various shapes; rarely bearing normal leaves, usually provided with spines or
barbed bristles (glochidia) which are generally considered to be modified leaves; spines and glochidia
develop from small cushion-like structures called areoles; in several genera, the stem assumes leaf-
like flattened form (e.g. Epiphyllum, Rhipsalis).
340 Plant Taxonomy
Leaves Simple, highly reduced or scale-like, or even absent in many genera, or flat and fleshy;
spines or tuft of bristles develop from areoles.
Inflorescence Flowers mostly solitary, showy and borne upon or near the areoles; only rarely corym-
bose or paniculate (Pereskia).
Flower Exceptionally beautiful, large, showy but only very rare (Mammillaria, Opuntia, Cereus,
Echinocereus, Schlumbergera); sessile; mostly bisexual; actinomorphic or nearly zygomorphic;
epigynous.
Perianth Perianth parts numerous (Fig. 21.64), showing a gradual transition from sepals to petals;
spirally arranged; sepals often petalloid; petals epigynous and present in several series; often fuse
to form a perianth tube or hypanthium.
Androecium Stamens numerous, in several series or groups, epipetalous or inserted at the base of
petals; anthers 2-celled, basifixed, introrse; longitudinally dehiscent.
Gynoecium Two to many carpels, syncarpous; ovary inferior; unilocular, numerous anatropous
ovules; parietal placentation, very rarely basal placentation; style 1 and simple; stigmas often as
many as the carpels.
Fruit and Seeds Fruit usually a many-seeded berry, often spiny, scaly, bristly or glochidiate. Seeds
with scanty or no endosperm; often arillate, with curved or straight embryo.
General Floral Formula ≈, , Pμ or (μ), Aμ , G (2 – μ).

• Cactaceae are mainly of ornamental value. Some commonly grown ornamental cacti include
species of Cereus, Cephalocereus, Echinocactus, Echinocereus, Mammillaria, Opuntia,
Rebutia and Stenocereus.
flower
Flowering Branch V.S. Flower Floral Diagram
Fig. 21.64 Opuntia dillenii Haw.

• Fruits of Nopalea cochenillifera, Lophophora williamsii and several Opuntia species are
edible.
• Stems of several cacti are used as emergency fodder.
• A red dye, cochineal, used in cosmetics, is obtained from several species of Opuntia.
• Some species of Cereus and Opuntia have medicinal properties.

Systematic position and relationships of family Cactaceae are highly controversial. Bentham and
Hooker treated Cactaceae under order Ficoidales and placed in between Passiflorales and Umbellales.
Hutchinson (1973) included it under order Cactales (placed in between Cucurbitales and Tiliales),
Cronquist (1981) included it under order Caryophyllales (placed in between Casuarinales and
Polygonales), and Thorne (1983) discussed it as the last family of order Chenopodiales.
Engler placed Cactaceae under order Opuntiales and opined this order to have been derived from
Parietales. Hutchinson considered that Cactaceae is allied to Cucurbitaceae while Maheshwari and
others suggested on the basis of the evidence of embryology and floral morphology that Cactaceae
show close affinities to the families included under Centrospermae or Caryophyllales. On the basis of
the recent discovery of the presence of the betalain pigments rather than anthocyanins, taxonomists
such as Cronquist (1981) and Jones and Luchsinger (1987) also support the placement of Cactaceae
under order Caryophyllales.
Britton and Rose (1919–1923), in their 4 volume works entitled “Cactaceae”, divided the family
into following three tribes:
1. Pereskieae (leaves broad, flat; glochids absent; flowers stalked), e.g. Pereskia.
2. Opuntieae (leaves, if present, more or less terete, small; glochids present; flowers sessile),
e.g. Opuntia.
3. Cacteae or Cereeae (leaves absent or rudimentary; glochids absent; flowers sessile), e.g.
Mammillaria, Cereus, Echinopsis, Epiphyllum, Rhipsalis, Schlumbergera.
21.50 UMBELLALES
1. Plants show simplification of their floral parts.
2. Flowers arranged in simple or compound umbel.
3. Flowers are usually bisexual, actinomorphic, pentamerous and epigynous.
4. Carpels reduced to 2, syncarpous, ovary inferior.
5. Each locule contains a single anatropous and pendulous ovule.
6. Style usually possesses a swollen base called stylopodium.
7. Fruit usually a dry schizocarp, called cremocarp, splitting into two mericarps.
Bentham and Hooker placed 3 families (Umbelliferae, Araliaceae and Cornaceae) under Umbellales.
Engler and Diels also divided order Umbelliflorae into same 3 families.
Only Umbelliferae and Araliaceae are treated in the present text.
342 Plant Taxonomy
21.51 UMBELLIFERAE OR APIACEAE CARROT FAMILY

Polypetalae, Calyciflorae, Umbellales.

Aromatic herbs; leaves compound and leaf base sheathing or pericladial; inflorescence simple or
compound umbel; inferior ovary; stylopodium present; fruit schizocarp.

Centella, Coriandrum, Daucus, Ferula, Foeniculum.

A family of about 275 genera and 3,000 species, Umbelliferae are cosmopolitan in distribution and
chiefly found in North Temperate Zone of the World. Over 50 of its genera and about 200 species
have been reported from India. Some of its larger genera with over 100 or more reported species
include Eryngium (230), Pimpinella (150), Bupleurum (150), Ferula (133), Peucedanum (120), and
Hydrocotyle (100).
The great Greek philosopher Socrates was put to death by using a weed of this family, Conium
maculatum (poison hemlock).

General Habit Mostly biennial or perennial herbs with stout stems and hollow internodes; some are
prostrate herbs (Centella, Hydrocotyle), few species of Bupleurum and Trachymene are somewhat
shruby; emit aromatic smell due to the presence of essential oil or oleoresin in all organs.
Leaves Alternate, generally pinnately compound or decompound, much divided, exstipulate, with
sheathing leaf bases; rarely leaves are entire and simple (Bupleurum, Centella, Hydrocotyle) or pal-
mately compound (Sanicula, Astrantia); bi- or tripinnately divided (Coriandrum, Foeniculum).
Inflorescence Usually a compound umbel subtended by an involucre of bracts; or simple umbel
(Astrantia, Hydrocotyle); or cymose head (Eryngium); compound umbels of majority of genera
are formed from partial umbels (each one often subtended by an involucre of bracteoles called
involucel).
Flower Usually bisexual, actinomorphic, pentamerous, epigynous; rarely unisexual because the last
umbellets to develop are sometimes staminate.
Calyx 5 sepals, free or united; very small and narrow, usually adnate to the ovary; valvate or rarely
imbricate (Astrantia).
Corolla 5 petals, polypetalous; valvate (Centella, Fig. 21.65) or rarely imbricate; petals alternate
with sepals; usually white or yellow; some deciduous, rarely absent; often bilobed; petal lobes of
peripheral flowers of Coriandrum are unequal.
Androecium 5 free stamens, alternate with the petals, inserted on

an epigynous disc; anthers dithecous, basi- or dorsifixed, introrse,
Gynoecium Bicarpellary, syncarpous, carpels medianly-placed;
ovary inferior, bilocular; each locule contains a single anatropous
and pendulous ovule; axile placentation; styles 2, often with a
thickened or swollen base (stylopodium) appearing like an epigy-
nous disc; stigmas 2.
Fruit and Seeds Fruit a dry schizocarp (called cremocarp) split-
ting into 2 mericarps, often ribbed. Two mericarps separate from
below upward from a stalk called carpophore. Each mericarp
contains a single seed. On the outer surface of each mericarp are
usually present 5 primary ridges, of which 2 lateral ridges are at
the edges where the splitting takes place. In the furrows between
the ridges are often present oil canals called vittae. Fruit structure
Fig. 21.65 Floral diagram
is an important diagnostic character in the taxonomy of this fam-
of Centella asiatica.
ily. Seeds with a minute embryo and copious endosperm.
General Floral Formula ≈, , K(5) or 5, C5, A5, G (2).

• Vegetables: Certain plants of Apiaceae, which provide us vegetables, which are either
eaten raw or cooked, include Apium graveolens (Celery), Daucus carota (Gajar or Carrot),
Pastinaca sativa (Parsnip) and Petroselinum crispum (Parseley).
• Condiments, Spices and Flavouring Agents: Mature fruits and seeds of several plants are
used as spices, condiments and for flavouring biscuits, breads, cheese, etc. Common spe-
cies are Anethum graveolens (Sowa or Dill), Apium graveolens (Celery seeds or Ajmud),
Carum carvi (Caraway or Shiajeera), Coriandrum sativum (Dhania or Coriander), Cuminum
cyminum (Zeera or Cumin), Foeniculum vulgare (Saunf or Fennel), Trachyspermum ammi
syn. Carum capticum (Ajwain), T. roxburghianum (Ajmud). Almost all these plants are
carminative, stimulant and used in stomach disorders as well as in many dishes in India,
China and several other countries.
• “Heeng”: ‘Heeng’ of commerce or Asafoetida is the dried latex obtained after making
incisions in the roots and rootstocks of Ferula assafoetida and F. narthex. Heeng is an
oleoresin and used as a condiment as well as in medicines. It stimulates respiratory as well
as nervous and intestinal systems.
• Poison: Conium maculatum (poison hemlock) is a deadly poison and was used to kill the
great Greek philosopher, Socrates. Cicuta virosa (water hemlock) is also poisonous.
• Medicinal Value: Centella asiatica (Brahmi Booti) is the king of medicinal plants. It is
used as a brain tonic and also in madness and leprosy. Ferula sumbule is used in curing
hysteria.
344 Plant Taxonomy
inflorescence
stigma
style
stylopodium
ovary
pedicel Fruit
Gynoecium
leaf
petal
stamen
stigma
style
stem
sepal
ovule
ovary
pedicel
L.S. Flower roots
Flowering Plant
Floral Diagram (zygomorphic flower) Floral Diagram (actinomorphic flower)
Fig. 21.66 Coriandrum sativum L.

• Ornamental Plants: Some of the ornamentals of Apiaceae include Angelica archangelica

(Angelica), Aegopodium podograria, Heracleum maxima and Trachymene caerulea.

Bentham and Hooker placed Umbelliferae under order Umbellales. Pollen characters of Umbelliferae
and Araliaceae show close affinities between these two families. However, Hutchinson opined that
Araliaceae is a primitive family and should be placed under Araliales of Lignosae while Umbelliferae
is a much advanced family of Herbaceae. In reduction of calyx and bicarpellary, syncarpous ovary
with 2 separate styles, Umbelliferae resembles Asteraceae (Compositae). Hutchinson opined that
Umbelliferae is derived from Saxifragales. However, Cronquist believed that Umbelliferae originated
from Sapindales.
Engler divided Umbelliferae into 3 subfamilies and 12 tribes as under:
I. Hydrocotyloideae: 1. Hydrocotyleae, 2. Mulineae.
II. Saniculoideae: 3. Saniculeae, 4. Legoiceae.
III. Apioideae: 5. Echinophoreae, 6. Scandiceae, 7. Coriandreae,
8. Smyrnieae, 9. Ammieae, 10. Peucedaneae,
11. Laserpitieae, 12. Dauceae.

Coriandrum sativum L. (Dhania or Coriander, Fig. 21.66)
Habit: Annual aromatic herb. Leaf: Alternate, exstipulate, decompound; upper leaves more finely dis-
sected; leaf base sheathing; unicostate reticulate. Inflorescence: Compound umbel. Flower: Bracteate
or ebracteate, complete, hermaphrodite, central flowers actinomorphic while peripheral flowers are
zygomorphic, pentamerous, epigynous. Calyx: 5 sepals, gamosepalous, free at the tips, valvate;
anterior sepals may be larger in outer peripheral flowers. Corolla: 5 petals, free, bilobed, valvate;
in central actinomorphic flowers the petal lobes are equal-sized; in peripheral zygomorphic flowers
one anterior petal has 2 large equal-sized lobes, two lateral petals have 2 unequal-sized lobes, and
two posterior petals have 2 equal-sized small lobes. Androecium: 5 stamens, alternipetalous, free,
dithecous, dorsifixed, introrse. Gynoecium: Bicarpellary, syncarpous, inferior, bilocular, one ovule in
each locule, axile placentation; styles 2, arise on a nectar-secreting disc called stylopodium; stigmas
2, capitate. Fruit: Cremocarp, splitting into 2 mericarps.
Floral Formulae:
(a) Peripheral flowers: , , K5 or (5), C5, A5, G (2).
(b) Central flowers: ≈, , K5 or (5), C5, A5, G (2).
21.52 ARALIACEAE ARALIA OR GINSENG FAMILY

Polypetalae, Calyciflorae, Umbellales.
346 Plant Taxonomy

Leaves alternate, often large and compound; usually umbellate inflorescence; actinomorphic, pen-
tamerous flowers; inferior ovary with one ovule in each locule; fruit berry or drupe.

Aralia, Helwingia, Panax.

A family of about 70 genera and over 800 species, Araliaceae are distributed chiefly in tropical
region, mainly in tropical America and Indomalaysia. In India, Helwingia himalaica occurs com-
monly in regions of Darjeeling while Panax fruticosum and P. pseudo-ginseng in Meghalaya and
Khasi Hills. Some other major genera along with their approximately reported species are Schefflera
(200), Aralia (35), Hedera (15, Fig. 21.67), Mackinlaya (12) and Panax (8). Members of Araliaceae
show close resemblances with that of Apiaceae.

General Habit Herbs, shrubs, sometimes vines or trees; some appear like palms; a few are root
climbers (e.g. Hedera).
Stem Solid, often prickly; In Hedera helix, the woody stem is evergreen, climber or ground-
carpeting and clothed densely by adhesive roots (Fig. 20.67).
Leaves Large-sized, compound, alternate, rarely opposite or whorled; glabrous; dark green; pal-
mately tri- or pentalobed (Hedera helix); stipulate, stipules usually modified as a membranous border
of petiolar base or linguliform.
Inflorescence Flowers small, forming umbels or heads, often grouped into compound inflorescences;
umbels sometimes racemose, corymbose paniculate or umbellate.
Flowers Small, bracteate, bisexual or unisexual (either on separate plants or occasionally male and
female flowers on the same plant); actinomorphic; usually epigynous and pentamerous; whitish or
greenish.
Calyx Usually 5, very small sepals, fused or adnate to ovary representing only small teeth-like
structures; cupuliform.
Corolla Usually 5 but sometimes 3–10 petals; free, caducous; often valvate.
Androecium Stamens 5, sometimes 3, generally equal to the number of petals and alternipetalous;
attached to the disc; anthers dithecous, dorsifixed, dehiscing longitudinally; anthers versatile in
Hedera helix (Fig. 21.67).
A nectariferous epigynous disc, covering top of the ovary and usually fused with the stylar bases,
is present.
Gynoecium Pentacarpellary, syncarpous, usually inferior ovary; rarely 1 to many fused carpels with
half-inferior or superior ovary; pentalocular, each locule with one pendulous and anatropous ovule;
axile placentation; styles as many as carpels, free or united; sometimes even absent, making the
stigmas then sessile.
stigma (sessile)
E
L.S. Inferior Ovary
leaf
A
Flowering Twig
C
adhesive
roots Inflorescence
stem
B
A Vegetative Shoot
H
stigma (sessile) A Fruit
locule
stamen
ovary ovule
petals ovary
wall
disc
G
F
T.S. Ovary A Stamen
D
A Flower
Fig. 21.67 Hedera helix.
Fruit Berry, rarely a drupe; with as many seeds as carpels.

Seed Seeds with small embryo and copious endosperm.
General Floral Formula Br, ≈, , K(5), C5, A5, G(5).
348 Plant Taxonomy

Araliaceae is not of much economic importance, except that of following major aspects:
• It provides some evergreen climbing vines (e.g. Hedera helix) and beautiful ornamental
shrubs and small trees belonging to several species of the genera like Aralia, Acanthopanax,
Schefflera, Fatsia, Kalopanax and Polyscias. Canary Island ivy (Hedera canariensis) is also
a well known plant of ornamental value.
• Chinese rice paper is obtained from the pith of rice-paper plant (Tetrapanax papyriferus).
• Ginseng roots, used in several medicinal preparations, are obtained from Panax ginseng and
P. quinquefolius. Ginseng is widely used in China as an stimulant and aphrodisiac.
• Species of Dizygotheca and Schefflera are cultivated indoors as pot plants. Several culti-
vars have been developed from Hedera helix showing wide range of variegation and leaf
shape.

Bentham and Hooker included three families under Umbellales, namely Cornaceae, Araliaceae and
Umbelliferae (Apiaceae). Hutchinson, however, discussed Cornaceae under order Cunoniales and
Araliaceae under order Araliales of division Lignosae, besides discussing Umbelliferae under order
Umbellales of division Herbaceae. According to him, Araliales have been derived from Cornaceae.
Hickey and King (1988) divided family Araliaceae into three tribes, viz. (1) Schefflereae
(corolla having valvate aestivation and petals with broad base), e.g. Hedera, Schefflera, (2) Aralieae
(corolla having imbricate aestivation and petals with broad base, e.g. Aralia, and (3) Mackinlayeae
(corolla having valvate aestivation and petals are shortly clawed).
Hallier recognised only 2 families under order Umbelliflorae, namely Cornaceae and Umbelliferae
and included all Araliaceae members under Umbelliferae.
21.53 GAMOPETALAE
The members of subclass Gamopetalae of Dicotyledons contain their flowers with partially or
completely fused petals of their corolla. It is divided into 3 series, viz. Inferae, Heteromerae and
Bicarpellatae.
Series Inferae is characterised by the presence of inferior ovary and contains 3 cohorts, viz.
Rubiales, Asterales and Campanales.
Series Heteromerae is characterised by the (i) presence of superior ovary; (ii) androecium of
one or two series, and (iii) presence of more than 2 carpels. It contains 3 cohorts, viz. Ericales,
Primulales and Ebenales.
Series Bicarpellatae contains (i) superior ovary, (ii) androecium of only one series, and (iii) only
2 carpels. It contains 4 cohorts, viz. Gentianales, Polemoniales, Personales, and Lamiales.
21.54 RUBIALES
1. Leaves are usually simple and opposite decussate.
2. Flowers are arranged in cymose inflorescences, generally dichasial cymes.
3. Flowers gamopetalous, usually actinomorphic but sometimes also zygomorphic.
4. Stamens epipetalous and inserted in the corolla tube.
5. Ovary bicarpellary, syncarpous, with 1 to many ovules in each locule; inferior ovary.
Bentham and Hooker included only two families in Rubiales. These are Rubiaceae and
Caprifoliaceae. Engler and Diels, however, also included 3 more families along with Rubiaceae and
Caprifoliaceae. These are Adoxaceae, Dipsacaceae and Valerianaceae.
Only Rubiaceae and Caprifoliaceae are treated in this text.
21.55 RUBIACEAE COFFEE FAMILY OR MADDER FAMILY

Gamopetalae, Inferae, Rubiales.

Herbs (mostly in north temperate regions), shrubs or trees; leaves stipulate, opposite or whorled;
flowers tetra- or pentamerous; stamens alternipetalous and as many as corolla lobes; usually bicarpel-
lary; ovary inferior.

Cinchona, Coffea, Gardenia, Ixora, Mussaenda, Oldenlandia, Rubia.

A family of about 500 genera and 6500 species, Rubiaceae are chiefly distributed in tropics, but
some are distributed in temperate or even arctic (Galium) regions. About 75 genera and 275 species
have been reported from India, chiefly in tropical and subtropical eastern Himalayas. Some of the
largely represented genera with their approximately reported species (Hickey and King, 1988) include
Psychotria (700), Galium (400), Ixora (400), Gardenia (250), Mussaenda (200), Asperula (200),
Cephaelis (180), Rondeletia (120), Coprosoma (90), Guettarda (80), Rubia (60), Coffea (40) and
Cinchona (40). Two most important plants of the family are Coffea arabica (coffee-yielding plant)
and species of Cinchona (quinine-yielding plant).

General Habit Mostly trees (Cinchona officinalis) or shrubs (Mussaenda luteola, Hamelia patens);
however, those in north temperate regions are mostly herbs (e.g. species of Galium, Oldenlandia);
sometimes hook climber (Uncaria) or vines.
350 Plant Taxonomy
Leaves Simple, entire; opposite decussate or whorled; stipulate, stipules usually interpetiolar and
rarely intrapetiolar, sometimes foliaceous (Galium) and become as large as the leaf blades so that
the leaves appear whorled; or sometimes reduced to glandular setae (Pentas).
Inflorescence Basically a dichasial cyme, dichasia sometimes grouped into globose heads (Adina,
Nauclea); rarely flowers solitary (Gardenia, Randia).
Flower Bracteate, sometimes bracteolate; bisexual; usually actinomorphic, rarely slightly zygomor-
phic; penta- or tetramerous; usually epigynous, only rarely perigynous (Synaptantha) or very rarely
hypogynous (Gaertnera).
Calyx 4 or 5 sepals, polysepalous, valvate; rarely greatly reduced sepals (Morinda); in Mussaenda
one of the sepals of some flowers becomes enlarged and bright coloured (Fig. 21.68).
Corolla 4 or 5 petals, gamopetalous; usually salverform, rotate, or infundibular; valvate (Ixora,
Mussaenda), twisted (Gardenia) or imbricate (Rondeletia).
Androecium 4 or 5 stamens, epipetalous, inserted in the corolla tube or at its mouth; alternate with
the corolla lobes; anthers 2-celled, basifixed, introrse, dehiscing longitudinally.
Gynoecium Usually bicarpellary, syncarpous, inferior ovary; bilocular, with 1 to many anatropous
ovules in each locule; axile placentation; style simple or bifid; stigma capitate; epigynous disc often
present; 5 carpels in Gardenia and Hamiltonia; ovary half-inferior in Synaptantha, or even rarely
superior in Gaertnera.
Fruit and Seeds Fruit a capsule (Cinchona, Oldenlandia), berry (Mussaenda), schizocarp (Galium),
or even drupe. Seeds with a small straight or curved embryo in endosperm; sometimes winged.
General Floral Formula Br, ≈, , K4 or 5, C (4)or (5), A4 or 5 , G (2).

• Ornamental Plants: Some of the common ornamental plants grown in gardens and road-
sides include Anthocephalus cadamba (Kadam), Cephalanthus occidentalis, Galium verum
(Ladies Bedstraw), Gardenia jasminoides (Jasmine), G. lucida, Hamelia patens, Hamiltonia
suaveolens (Padera), Ixora arborea, I. coccinia, Morinda tinctoria, Mussaenda frondosa, M.
luteola, Pentas lanceolata and Rondeletia.
• Coffee: Coffee, the most popular nonalcoholic, caffeine-containing beverage of the world,
comes mainly from roasted and powdered seeds of Coffea arabica. Other coffee-producing
species are C. liberica and C. robusta.
• Quinine: This famous drug, used throughout the world in the treatment of malaria, comes
from the bark of several species of Cinchona, such as C. calisaya, C. ledgeriana and
C. officinalis.
• Other Medicinal Plants: (i) Cephaelis ipecacuanha (Ipecac) roots are used against amoe-
bic dysentery and pyorrhoea, (ii) Gardenia gummifera provides gum used as a carminative
stimulant, (iii) Randia tinctoria fruit pulp is emetic and anthelmintic, (iv) Paederia foetida
is used in indigestion and stomach troubles.
petal flowers
anther
lobe
connective
stamen corolla
filament
tube
Stamen
bract
stigma
stipule
style
sepal ovary wall

modified locule
leaf
ovule sepal ovule
placenta
stem
ovary
pedicel
L.S. Flower
T.S. Ovary
Flowering Branch
Floral Diagram (actinomorphic flower) Floral Diagram (zygomorphic flower)
Fig. 21.68 Mussaenda luteola Dilile.

352 Plant Taxonomy
• Dyes: Roots of Rubia tinctoria (Madder) contain dye alizarin and purpurin, while that of
several species of Morinda contain red (M. tinctoria, M. bracteata) and yellow (M. citridora)
dye. Red dye is also obtained from the root bark of Oldenlandia umbellata.
• Timber: Useful timber is obtained from several species of this family, such as Adina cor-
difolia (Haldu), Anthocephalus cadamba, Ixora ferrea, Mitragyna parviflora and Randia
spinosa.

Bentham and Hooker treated Rubiaceae under order Rubiales while Takhtajan (1969) and Thorne
(1983) discussed it under order Gentianales.
Affinities of Rubiaceae are controversial. Similar type of alkaloids bring Rubiaceae close to
Loganiaceae. But in Loganiaceae the ovary is superior while it is inferior in Rubiaceae. In possessing
opposite leaves, cymose inflorescence and inferior ovary Rubiaceae comes closer to Caprifoliaceae.
However, leaves are exstipulate in Caprifoliaceae. In possessing epigynous flowers, cymose inflo-
rescence, epigynous disc and bicarpellate ovary, Rubiaceae comes closer to Umbelliferae and
Cornaceae.

Mussaenda luteola Dilile (vern. Bedina, Fig. 21.68)
Habit: An ornamental, perennial shrub. Leaf: Opposite decussate, simple; stipulate, stipules inter-
petiolar; subsessile, acute; unicostate reticulate. Inflorescence: Dichasial cyme. Flower: Bracteate,
pedicellate, complete, hermaphrodite; younger flowers actinomorphic, older flowers zygomorphic;
pentamerous, epigynous. Calyx: 5 sepals, free but sometimes fused, valvate; in zygomorphic flow-
ers one of the sepals is modified into a large, yellow, leaf-like structure. Corolla: 5 petals, fused,
valvate, hypocrateriform, throat and mouth of corolla hairy. Androecium: 5 stamens, polyandrous,
epipetalous, alternipetalous; dithecous, basi- or dorsifixed, introrse. Gynoecium: Bicarpellary, syn-
carpous, inferior, bilocular, with many ovules on T-shaped placentae; axile placentation; style long;
stigma bilobed. Fruit: Berry.
Floral Formula: Br, ≈ or , , K5, C (5), A5, G (2).
21.56 CAPRIFOLIACEAE HONEYSUCKLE FAMILY

Gamopetalae, Inferae, Rubiales.

Shurbs or lianas, rarely herbs; opposite; exstipulate leaves; flower tetra- or pentamerous, epigynous;
calyx fused to the ovary; inferior ovary; multicarpellate gynoecium.

Lonicera, Viburnum, Wiegela, Abelia.

Caprifoliaceae is a family of about 15 genera and more than 530 species, distributed mostly in north-
temperate regions and tropical mountains of the world. Some of the larger genera of the family are
Lonicera (200, honeysuckle) distributed mainly in the North America and Eurasia, Viburnum (200)
distributed in temperate regions and subtropics, Sambucus (40) showing a cosmopolitan distribution,
Abelia (30) found chiefly in Himalayas to east Asia and Mexico, and Symphoricarpos (18) distributed
in North America and China.
Sweet nectar, obtained from the flowers of Lonicera (honeysuckle; Fig. 21.69), is enjoyed by mil-
lions of children throughout the world.

General Habit Shrubs, sometimes lianas (Lonicera sp.); only rarely herbs (Triosteum); Lonicera
periclymenum (Fig. 21.69), a woody climber, reaches upto 6 m.
Leaves Simple but sometimes pinnately compound (Sambucus); opposite; usually exstipulate but
sometimes stipulate in Sambucus, stipules reduced to nectariferous glands in m; margin
entire but occasionally lobed in some members; ovate to elliptical in Lonicera periclymenum
(Fig. 21.69).
Inflorescence Cymose or its modifications.
Flowers Usually bisexual; actinomorphic or sometimes zygomorphic (Lonicera periclymenum,
Fig. 21.69); in some members, the 2-flowered cymes are bracteolate; in some species of Viburnum,
the flowers are sterile or neutral.
Calyx Sepals 4 or 5, gamosepalous; usually fused to the ovary; sometimes polysepalous.
Corolla Petals 5, sometimes 4, gamopetalous, imbricate; sometimes bilabiate or variable in form;
epigynous.
Androecium Stamen 5, epipetalous, attached on corolla tubes; alternipetalous; sometimes stamens
4 due to suppression of posterior stamen as in Linnaea; anthers usually introrse as well as extrorse
as in Sambucus.
Gynoecium Compound pistil of usually 3–5 carpels; syncarpous; ovary inferior; locules as many as
carpels, each containing 1 or more pendulous ovules, axile placentation; style terminal, often slender;
stigmas capitate and as many as carpels, distinct or united.
Fruit A berry or drupe; sometimes an achene or capsule.
Seeds With abundant fleshy endosperm, usually each with a small and straight embryo.
General Floral Formula Brl, ≈, , K(5), C (5), A5, G (3 – 5).

Caprifoliaceae is important for a number of ornamental shrubs, vines and edible fruits.
• Sweet nectar obtained from the flowers of honeysuckle (Lonicera) has been enjoyed by the
children of countless generations.
354 Plant Taxonomy
unopened glandular
flowers flower corolla
glandular
calyx
locule
ovary
ovules
D bract
leaf
L.S. Inferior Ovary
A
stem
locule
A. Flowering Branch
ovules
(2)
anthers
(versatile)
E
T.S. Ovary
upper lip
(4-lobed)
filament
stamens
C
A Stamen stigma
(Upper Part) (trilobed)
corolla tube
glandular hairs
style
B
bilipped corolla
L.S. Corolla Tube
lower lip
Fig. 21.69 Lonicera periclymenum L.
• Fruits of elderberry (Sambucus nigra) are used to make elderberry wine.

• Over 200 species of several genera (Viburnum, Lonicera, Abelia, Leycesteria, Kolkwitzia,
Symphoricarpos, Diervilla and Linnaea are used as showy ornamental shrubs.
• Fruits of Viburnum trilobum (highbush cranberry) are edible and also used in preparing
jelly.
• Lonicera japonica is a noxicus troublesome weed and should be taken with all care.

Treated under order Rubiales of series Inferae of Gamopetalae by Bentham and Hooker, honey-
suckle family (Caprifoliaceae) has been included under order Dipsacales of subclass Asteridae by
Cronquist (1981) along with three other families, viz. Adoxaceae, Valerianaceae and Dipsacaceae.
Most phylogenists have treated the monotypic genus Adoxa in a separate family Adoxaceae. The
genus Sambucus, sometimes placed in a separate family Sambucaceae, should be treated only under
Caprifoliaceae because of its several similarities with the family, such as (i) pinnate leaves, and (ii)
extrorse anthers.
21.57 ASTERALES
1. Herbs, shrubs or vines; only rarely trees.
2. Inflorescence an involucrate head or capitulum.
3. Stamens epipetalous, syngenesious or synandrous or free.
4. Bicarpellary, syncarpous; ovary inferior, unilocular, one ovule in the locule.
5. Placentation basal.
Bentham and Hooker included Compositae and 3 more families under Asterales. Engler and Diels,
however, included Compositae under order Campanulatae along with 5 more families (Campanulaceae,
Brunoniaceae, Calyceraceae, Goodeniaceae and Stylidaceae).
Only Compositae (=Asteraceae) has been discussed in this text. Campanulaceae has been treated
under Campanales and has been discussed with this order elsewhere in this text.
21.58 COMPOSITAE OR ASTERACEAE SUNFLOWER OR ASTER FAMILY

Gamopetalae, Inferae, Asterales.

Herbs, vines, or shrubs, rarely trees; inflorescence an involucrate head or capitulum; pappus often
present; stamens syngenesious; carpels 2, united; ovary inferior; fruit cypsela.

Ageratum, Aster, Chrysanthemum, Dahlia, Eclipta, Helianthus, Launaea, Sonchus, Vernonia.
356 Plant Taxonomy

Compositae is one of the largest families of flowering plants, represented by about 1100 genera and
over 20,000 species (Jones and Luchsinger, 1987), which are worldwide in distribution. Radford (1986)
has mentioned the number of genera to be “One thousand to 2000”. About 140 of its genera and
over 700 species have been reported from India. Some of the largely represented genera with their
approximately known number of species (Jones and Luchsinger, 1987; Hickey and King, 1988) in
brackets include Senecio (2000+), Eupatorium (1200), Vernonia (1000), Hieracium (1000), Centaurea
(600), Aster (500), Helichrysum (500), Cousinia (400), Artemisia (400), Baccharis (400), Mikania
(250), Bidens (230), Crepis (200), Inula (200), Achillea (200), Gnaphalium (200), Chrysanthemum
(200), Anthemis (200), Erigeron (200), Cirsium (150), Ligularia (150), Helianthus (110), Lactuca
(100), Hypochoeris (100), and Carduus (100).

General Habit Usually annual or perennial herbs, but some are shrubs and only a few are trees
(Leucomeris, Vernonia arborea); Senecio, the largest genus of the family includes herbs (S. vulgaris),
shrubs (S. magnificus), trees (S. cruentus) and even climbers (S. scandens); majority of the temper-
ate zone Compositae are herbaceous; sap watery or milky; usually with rhizomes, stolons, tubers,
or fleshy roots; plants often with spiny or various types of vestiture or surface coverings, such as
tomentose, lanate, pannose, strigose, etc.; few are xerophytes, e.g. Hoplophyllum.
Leaves Usually alternate, sometimes opposite (Dahlia, Helianthus), rarely whorled (Eupatorium);
mostly simple, sometimes needle-like or reduced to scales; margins often pinnately or palmately
lobed or divided; only rarely truly compound; exstipulate; frequently in basal rosettes; leaves usually
with oil passages; some contain latex (Lactuca, Crepis).
Inflorescence The primary inflorescence is a head or capitulum with many (rarely 1) flowers (called
florets) borne on a conical, flat, concave, or convex receptacle (Fig. 21.70); each floret is often sub-
tended by a receptacular bract called pale or chaff; the receptacle is often subtended by an involucre
of bracts called phyllaries; bracts or phyllaries are of various shapes, sizes, and textures, and present
in one or more series. Usually the heads or capitula are arranged in various ways, such as racemes,
panicles, corymbs, or compound heads; in Echinops, the heads are compound, but each smaller head
contains only one flower. The capitulum may be heterogamous or homogamous.
Flower Flowers are actinomorphic or zygomorphic; bisexual, or unisexual, or even neutral (both
stamens and carpels are aborted); pentamerous; epigynous.
In heterogamous capitulum inflorescence (e.g. Helianthus, Fig. 21.71), the central florets are disc
florets which are bisexual and actinomorphic; and peripheral florets are ray florets which are ligulate,
generally female or neutral and zygomorphic.
In homogamous capitulum inflorescence, all florets are similar, usually bisexual, actinomorphic
or zygomorphic, rarely all unisexual.
Undermentioned are the five basic plans of florets found within the heads or capitula of
Compositae:
1. Tubular or disc florets, corolla regular, florets perfect.
2. Tubular or disc florets, corolla regular, florets pistillate.
stigma
style
ray floret
pappus
disc floret involucral bract

(phyllary)
ligule (formed from

5 united petals)
receptacle
stigma
style
style
united anthers Inflorescence (head) united
with Left half Section anthers
ray floret
disc floret
corolla
corolla tube
involucral bract
(phyllary) chaffy bract
pappus
pappus
inferior ovary receptacle
peduncle
ovary
Ray Floret V.S. Head of Compositae
(diagrammatic) Disc Floret
Fig. 21.70 Inflorescence and florets of Compositae.
3. Ligulate or ray florets, zygomorphic, pistillate.

4. Ligulate or ray florets, zygomorphic, sterile.
5. Ligulate or ray florets, zygomorphic, perfect.
Calyx Represented by pappus of bristles, awns, or scales; or absent; epigynous.
Corolla 5 petals, gamopetalous, valvate; represented by 3 basic types: (i) 5-lobed and tubular,
(ii) ligulate with 3 to 5 teeth, (iii) bilabiate with 3 lobes in the upper lip and 2 lobes in the lower
lip. In Mutisia, all florets are bilabiate while in Lactuca and Cichorium all florets are ligulate.
Androecium 5 stamens, epipetalous, alternating with corolla lobes; anthers nearly always united
into a tube around the style and filaments free i.e. syngenesious; dithecous, connective often
358 Plant Taxonomy
stigma flowers
style
bract
petal
stamen
thalamus
L.S. Head
stigma
sepal style
inflorescence
bract
ovary ovule
ovule bract ovary
L.S. Disc Floret L.S. Ray Floret

stigma
style
leaf
sepal
bract
stem
Disc Floret Ray Floret
Flowering Branch
Floral Diagram (disc floret)

Floral Diagram (ray floret)
Fig. 21.71 Helianthus annuus L.
prolonged, introrse, longitudinally dehiscent; in Silybium the stamens unite by anthers as well as
by filaments.
Gynoecium Bicarpellary, syncarpous; ovary inferior; unilocular; one ovule, anatropous; basal pla-
centation; style 1; usually 2 branched stigma of diverse forms; an epigynous nectar-secreting disc is
also present at the base of the corolla tube surrounding the style.
Fruit and Seeds Fruit a cypsela, often crowned by the persistent pappus, sometimes enclosed by
persistent bracts. Seed is 1 per fruit; with large and straight embryo and no endosperm.
Pollination The pollination mechanism is entomophilous and noteworthy in Compositae. A single
insect may pollinate numerous flowers of a capitulum in single visit. Insects are attracted by the
conspicuous ligulate ray florets of the capitulum. This also benefits the often inconspicuous disc
florets. Ray florets also protect the nectar from rain. Cross pollination is promoted also by the
protandrous condition of the androecium. In case of failure of cross-pollination, self-pollination may
also take place.
General Floral Formula of a Bisexual Flower or ≈, , Kpappus, C (5), A5, G (2).
21.58.6 Economic Importance1

• Ornamental Plants: Well-known ornamental species of the family include Aster amellus,
A. grandiflorus, Calendula officinalis, Centaurea moschata, Chrysanthemum carinatum, C.
coronarium, C. indicum, Coreopsis grandiflora, Cosmos bipinnatus, Dahlia excelsa, D. pin-
nata, Dimorphotheca sinuata, Gaillardia pulchella, Gynura aurantiaca, Helianthus annuus
(Sunflower or Surajmukhi), Helichrysum petiolatum, Tagetus patula (French Marigold or
Genda), Zinnia elegans and Z. linearis.
• Disease-Causing Plants: (i) Ambrosia artemissifolia (Rugweed)—pollen cause hay fever,
(ii) Eupatorium urticaefolium causes milk sickness in animals, (iii) Pyrethrum hysterophorus
(carrot grass)—pollen cause skin allergy.
• Medicinal plants: (i) Anthemis nobilis dried capitula are used against dyspesia, (ii)
Artemisia cina flower heads provide the drug santonin, an antidote against intestinal worms,
(iii) Blumea balsamifera leaves are used against excitement and insomnia while B. lacera
leaves are given to cure cholera, (iv) Grindelia camporum heads are used in bronchitis and
whooping cough, (v) Inula helium leaves are effective against tuberculosis, (vi) Lactuca
virosa leaves have sedative properties, (vii) Sphaeranthus indicus capitula are used for
curing stomach ache and piles, (viii) Spilanthes paniculata capitula are chewed to relieve
toothache, (ix) Tanacetum vulgare leaves are effective in chronic ulcers and rheumatism,
(x) Taraxacum officinale roots and rhizome provide the drug “taraxacum”, used as a mild
laxative, and (xi) Tussilago farfara leaves are used in asthama, cough and colds.
• Insecticides: Powdered dry capitula of Centrantherum anthelmenticum, Chrysanthemum
coccineum and Pyrethrum cinerariifolium are used in preparing insecticides.
• Edible Products: (i) Tuberous roots of Cynara scolymus, Dahlia tuberosa, Helianthus
tuberosus (Jerusalem artichoke) and Tragopogon porrifolius (vegetable oyster) are edible and
used as food by man, (ii) Lactuca sativa (Salad or Garden lettuce) leaves are eaten raw as
salad, (iii) roasted seeds of Taraxacum officinale are used in place of coffee.
• Oils: (i) Oil obtained from seeds of Carthamus tinctorius (safflower or Kusum) is used
for manufacture of soaps, paints, varnishes etc., (ii) hair-oil, ‘Bhrangraj’, is obtained from
1
Plants mentioned alphabetically in the subheadings.
360 Plant Taxonomy
the leaves of Eclipta alba, (iii) fatty oil, obtained from the seeds of Helianthus annuus, is
edible, (iv) a strong, aromatic essential oil, used in perfumery, is obtained from the seeds of
Tagetus patula (French Marigold), (v) oil from the seeds of Xanthium strumarium is used
as an illuminant.
• Dye: A red dye safflower, obtained from Carthamus tinctorius, is used for colouring
candles, butter, and several liquors.

Compositae is usually divided into 13 tribes. These tribes, along with one common example of each of
them, are (1) Heliantheae (Helianthus), (2) Astereae (Aster), (3) Anthemideae (Chrysanthemum), (4)
Arctotideae (Arctotis), (5) Inuleae (Inula), (6) Senecioneae (Senecio), (7) Calenduleae (Calendula),
(8) Eupatorieae (Eupatorium), (9) Vernonieae (Vernonia), (10) Cynareae (Carthamus), (11) Mutisieae
(Mutisia), (12) Liabeae (Liabum), and (13) Lactuceae (Lactuca).
In possessing pentamerous flowers, syngenesious anthers and inferior ovary, Compositae is related
to Campanulaceae and Goodeniaceae. In all these three families the ovary is usually bicarpellary.
In possessing inferior bilocular ovary, Compositae also comes near to Rubiaceae and Stylidiaceae.
The characters, such as dense inflorescence and inferior ovary, also trace the affinity of Compositae
with Dipsacaceae and Valerianaceae. It is presumed that Compositae originated from Rubiaceae.
21.58.8 Whether Compositae Occupy Highest Position Among Angiosperms?

Undermentioned are some advanced characters, because of which some workers regard Compositae
to be the most highly evolved taxon among angiosperms:
1. Cosmopolitan distribution from arctic to tropical zone.
2. Herbaceous habit.
3. Capitulum inflorescence.
4. Presence of large number of flowers in a small space which makes it possible to pollinate
several flowers easily by a single insect.
5. Presence of both actinomorphic and zygomorphic flowers.
6. Presence of both unisexual as well as bisexual flowers.
7. Presence of calyx in the form of pappus.
8. Gamopetalous corolla.
9. Syngenesious anthers.
10. Protandry of androecium.
11. Reduction of carpels to 2 and number of ovules to only one.
12. Presence of inferior ovary and basal placentation.
13. Cypsela fruit.
14. Elaborate mechanism of dispersal.

1. Ageratum conyzoides L. (Goat weed, Fig. 21.72)
Habit: Annual herb. Leaf: Simple, opposite, lower leaves alternate; exstipulate, ovate, serrate, acute,
hairy; unicostate reticulate. Inflorescence: Compound capitulum; different capitula arranged in
corymbose manner; inflorescence homogamous with all flowers tubular; outside each capitulum is
present involucre of bracts. Flower: Complete, actinomorphic, hermaphrodite, pentamerous, epigy-
nous. Calyx: 5 sepals, polysepalous; valvate; reduced to long scaly structures. Corolla: 5 petals,
gamopetalous; valvate; tubular. Androecium: 5 stamens, epipetalous, syngenesious; filaments short;
stigma
stigma
petal
stamen
style
style
inflorescence
leaf
ovary ovary
L.S. Flower stem Gynoecium
Flowering Branch
anther
lobe
connective
filament
Stamen
Floral Diagram
Fig. 21.72 Ageratum conyzoides L.

362 Plant Taxonomy
anthers dithecous, basifixed, introrse. Gynoecium: Bicarpellary, syncarpous, inferior ovary, unilocu-
lar, one ovule in the locule, basal placentation; style long; stigma bifid. Fruit: Cypsela.
Floral Formula: ≈, , K5 (pappus), C (5), A5, G (2).
21.59 CAMPANALES
1.
Flowers typically pentamerous with 5 sepals, 5 petals and 5 stamens.
2.
Corolla gamopetalous.
3.
Stamens usually in a single whorl, anthers are dithecous and often coherent to connate.
4.
Gynoecium is bi- to pentacarpellary, syncarpous.
5.
Ovary inferior; often unilocular but sometimes bi- to multilocular; when unilocular, the locule
contains a single ovule.
Cohort Campanales includes Campanulaceae and 3 more orders (=families) according to
Bentham and Hooker. Engler and Diels, however, placed 6 families in the order Campanulatae, viz.
Campanulaceae, Brunoniaceae, Calyceraceae, Goodeniaceae, Stylidaceae and Compositae.
Only Campanulaceae of Campanales has been treated here. Compositae of order Campanulatae
(according to Engler and Diels) has been discussed under order Asterales elsewhere in this text.
21.60 CAMPANULACEAE BELLFLOWER FAMILY

Gamopetalae, Inferae, Campanales.

Largely herbaceous, rarely shrubs or trees; leaves simple, alternate and exstipulate; flowers showy,
bisexual, epigynous, pentamerous; stamens frequently united, epipetalous and show connation of
their anthers or filaments; ovary inferior, with axile placentation.

Campanula, Lobelia, Centropogon, Cyphia.

A family of about 70 genera and over 2000 species, Campanulaceae are widely distributed in tem-
perate and sub-tropical regions of the world, specially in tropical mountains. The chief genera of the
family are Campanula (300), Lobelia (235), Centropogon (230), Siphocampylus (215), Wahlenbergia
(150), Cyphia (50), Cyanea (50), Phyteuma (40) and Lightfootia (40).
Several taxonomists treat Campanulaceae as a family distinct from Lobeliaceae. However, major-
ity of phylogenists, except Hutchinson, treat Campanulaceae comprising of 3 subfamilies (namely,
Campanuloideae, Cyphioideae and Lobelioideae, as has also been treated here in this text.

General Habit Annual or mostly perennial herbs or subshrubs, rarely trees (e.g. Clermontia); contain
milky or watery sap.
Leaves Simple, alternate or rarely opposite; exstipulate; contain latex; in Siphocampylus the leaves
are whorled; with long petiole in Campanula rotundifolia (Fig. 21.73).
Inflorescence Usually racemose, sometimes cymose as in Canarina; if cymose, it may be a deter-
minate monochasial or dichasial cyme; sometimes the flowers are arranged in involucral heads or
they are solitary axillary (Campanula rotundifolia, Fig. 21.73).
Flower Bracteate, often bracteolate, complete, hermaphrodite, actinomorphic or zygomorphic, pen-
tamerous, epigynous or rarely hypogynous (e.g. Cyananthus); blue, regular and bell-shaped in
Campanula rotundifolia (Fig. 21.73).
Calyx Sepals 5, rarely 3–10, fused with the ovary; valvate or imbricate.
Corolla Petals 5, gamopetalous, valvate; campanulate or tubular or bilabiate; in bilabiate flowers, the
corolla is zygomorphic and split down on one side; petal lobes are 6 in Canarina while only 3–4
in Edrianthus; instead of gamopetalous, the corolla is polypetalous in Phyteuma and even absent in
some species of Specularia; often blue or violet.
Androecium Stamens 5 or as many as corolla lobes, alternipetalous, distinct or variously connate;
bases of filaments often expand and form a dome-shaped chamber over epigynous disc; epipetalous
or distinct; anthers dithecous, distinct or connate, introrse.
Gynoecium Bi- to pentacarpellary, syncarpous; inferior; only rarely half-inferior in some species
of Lobelia and Edrianthus or even superior as in Cyananthus; usually two to many locules, with
many ovules in each locule on axile placentation; style slender and simple; stigmas usually 2–5 or
as many as carpels.
Fruits A capsule showing different ways of dehiscence in different genera; sometimes a berry (e.g.
Centropogon).
Seeds With fleshy abundant endosperm and small and straight embryo.
General Floral Formula Br, Brl, ≈ or , , K(5), C (5), A5 or A(5), G (2 – 5).

• Several members of Campanulaceae, with their showy flowers, are grown as garden orna-
mentals. Chief among them are species, hybrids and cultivars of Campanula (bell flower),
Codonopsis, Edrianthus, Lobelia, Phyteuma, Platycodon, Specularia and Sympyandra.
• The drug “Lobelia” is obtained from the dried leaves and tops of Indian tobacco (Lobelia
inflata), a small annual with blue flowers. The plant, as such, is poisonous and cultivated in
Kerala, Nilgiris and Darjeeling. The drug is used as an expectorant, antispasmodic, emetic
and also in chronic bronchitis and asthma.

Most taxonomists as well as phylogenists divide family Campanulaceae into three sub-families as
under:
364 Plant Taxonomy
petals
stamen stigma
stigma
style
E corolla
Reproductive Organs tube
sepal
B
leaf A Bisexual Flower anther
flower
filament
hairs
D
A Stamen
expanded base of
A filament
disc
ovules
A Flowering Plant locule
petal ovary
F
stigma L.S. Inferior Ovary
style
stamens locule
ovules
sepal
G H
T.S. Ovary
A Fruit
C
L.S. Flower Showing
Stamens and Style
Fig. 21.73 Campanula rotundifolia L.

(1) Campanuloideae, having actinomorphic and only rarely zygomorphic flowers with usually free
anthers, e.g. Campanula, Phyteuma, Jasione.
(2) Cyphioideae, having zygomorphic flowers with their stamens having their filaments united and
anthers free, e.g. Cyphia, Nemacladus.
(3) Lobelioideae, having zygomorphic flowers and their stamens having their anthers united, e.g.
Lobelia, Centropogon.
Some taxonomists, however, believe that members of Lobelioideae should be treated under inde-
pendent family Lobeliaceae due to several striking differences between Lobelia and Campanula.
21.61 ERICALES
1. Flowers generally actinomorphic, hypogynous and pentamerous.
2. Petals generally basally connate or sometimes distinct and free.
3. Stamens obdiplostemonous and usually inserted at the edge of a hypogynous nectariferous
disc.
4. Gynoecium usually pentacarpellary, syncarpous with superior ovary.
5. Numerous anatropous ovules in each locule; axile placentation.
6. Seeds very minute with copious endosperm.
Cohort Ericales, according to Bentham and Hooker, contains Ericaceae and 5 more families. Engler
and Diels, however, divided order Ericales into 2 suborders and 4 families i.e. suborder Ericineae
with 3 families (Ericaceae, Clethraceae, and Pyrolaceae) and suborder Epacridineae with only 1
family Epacridaceae.
Only Ericaceae is discussed in this text.
21.62 ERICACEAE HEATH FAMILY

Dicotyledons, Gamopetalae, Heteromerae, Ericales.

Predominantly shrubs; leaves alternate; flowers urceolate or campanulate; the stamens usually
distinct, mostly twice as many as of corolla lobes, developing from a nectariferous disc and only
rarely adnate to petals; anthers usually opening by apical pores; ovary typically tetralocular to
multilocular.
366 Plant Taxonomy

Rhododendron hookerii, Gaultheria fragrantissima, Lyonia ovalifolia.

Represented by about 125 genera and 3500 species (Jones and Luchsinger, 1987), members of
Ericaceae are cosmopolitan in their distribution. They are, however, almost absent from desert areas
and from Australasia where it is replaced by members of the allied family Epacridaceae (Hickey
and King, 1988). Ericaceae are mainly confined in the tropical regions, especially in acidic soils on
peat, swamps and woodlands. They are mainly associated with endotrophic mycorrhiza.
Major genera of the family with their approximately known number of species in parenthesis are
Rhododendron (850), Erica (600, heath), Vaccinium (450, blueberry or cranberry), Arctostaphylos
(70, manzanita) and Gaylussacia (50, huckleberry). Several handsome cultivars have been produced
by horticulturists by crossing various species of Rhododendron.

General Habit Woody shrubs, rarely small trees (Arbutus), or trailing or scrambling vines attaining
a length upto 20 metres.
Leaves Simple, usually alternate and evergreen or coriaceous; sometimes opposite or whorled;
exstipulate; entire, elliplical to oblong (Rhododendron ponticum, Fig. 21.74); sometimes linear and
in whorls of 4 (Erica herbacea).
Inflorescence Flowers solitary in axils or in axillary or terminal clusters; racemose racemes or
panicles; flowers form terminal umbel-like racemes in Rhododendron ponticum (Fig. 21.74) or leafy
and one-sided raceme in Erica herbacea.
Flowers Bracteate, bracteolate, usually bisexual (Epigaea) and actinomorphic but slightly zygomor-
phic as in Rhododendron ponticum; often showing wide range of colours.
Calyx Usually 4 to 7 sepals, generally gamosepalous but sometimes polysepalous; usually
persistent.
Corolla 4 to 7 petals, gamopetalous but sometimes distinct petals; urceolate, campanulate or salver-
shaped; convolute or imbricate aestivation.
Androecium Stamens as many or generally twice as many as petals developing from the base of
a nectar-secreting disc; disc is hypogynous in some members (e.g. Rhododendron) and epigynous
in Gaultheria; stamens free or fused at the base; obdiplostemonous; anthers dithecous, frequently
appendaged; anthers usually opening by apical pores; introrse.
Gynoecium Usually 4 to 5 carpels, sometimes upto 10 carpels, syncarpous; usually superior ovary,
with 4 to 5 locules, locules generally opposite the corolla lobes; numerous anatropous ovules; axile
placentation; style simple, usually 1, conical to filiform; stigma usually capitate; sometimes ovary
inferior (e.g. Vaccinium).
Fruit A capsule, berry or drupe.
Seed Usually small with cylindrical embryo and copious or fleshy endosperm.
General Floral Formula Br, Brl, ≈, , K(4 – 7), C (4 – 7), A4 – 7 or 8 – 14, G (4 – 5).
winter bud
stigma petals
style
scaly
leaves
stamens
(10)
D
A Flower
A
A Large Terminal Winter Bud
bract
terminal
pore
pedicel
H C
F T.S. Ovary
A Bract
A Stamen
style
ovules
sepal
ovary pedicel
disc
bract
calyx receptacle
disc
G B ovary
L.S. Ovary A Floral Bud E
Ovary
Fig. 21.74 Rhododendron ponticum.

From the point of view of economic importance, heath-family is important for its ornamentals
and for blueberries and cranberries.
• Ornamental shrubs are obtained from several species of Erica, Rhododendron, Kalmia,
Leucothoe and Pieris. Several beautiful hybrids and cultivars of horticultural value are
obtained from Erica (Heath), Rhododendron, Arbutus, Pernettya and Gaultheria. They are
commonly grown in gardens and glasshouses.
368 Plant Taxonomy
• The fruits of several species of Vaccinium are edible being variously known as cranberries
(V. macrocarpon), blueberries, bilberries and cowberries.
• Huckleberry, an edible fruit, comes from the genus Gaylussacia.
• Some of the suffrutescent perennials or subshrubs are Mayflower (Epigaea), bearberry
(Arctostaphylos uva-ursi) and wintergreen (Gaultheria procumbens).
• Foliage of Gaultheria shallon is sold in the market as “lemon leaf”.
• The so-called “briar-pipes” are obtained from the burls of Erica arborea.

Some taxonomists divide family Ericaceae, as circumscribed above, into two independent families,
viz. Ericaceae and Vacciniaceae, with the former having superior ovary and capsular fruit and the
latter possessing partially or completely inferior ovary and berry-type of fruit. Engler and Diels
divided Ericaceae into four subfamilies, namely Rhododendroideae, Arbutoideae, Vaccinioideae and
Ericoideae. Hickey and King (1988) divided Ericaceae into four subfamilies as under:
1. Rhododendroideae Possessing winter buds with scales; petals caducous; stamens without any
appendages, fruit septicidal capsule, and winged seeds, e.g. Rhododendron.
2. Ericoideae Possessing no scales or winter buds; persistent corolla lobes; leaves needle-like; sta-
mens with appendages; fruit loculicidal capsule, or nut, and; seeds with no wings, e.g. Erica.
3. Vaccinioideae Possessing winter buds with scales; leaves broad, flat and not needle-like; petals
caducous; stamens with appendages; inferior ovary; fruit a capsule, berry or drupe, and; seeds with
no wings; e.g. Vaccinium.
4. Epigaeoideae Dioecious plants; leaves cordate; stamens with no appendages; expanded stigma
with 5 lobes; e.g. Epigaea.
Bentham and Hooker treated Ericaceae under order Ericales of series Heteromerae of Gamopetalae
while Engler and Prantl discussed Ericales under subclass Metachlamydeae of Dicotyledoneae.
Hutchinson discussed order Ericales under Division Lignosae of subphylum Dicotyledones while
Takhtajan (1980) treated order Ericales under superorder Ericanae of subclass Dilleniidae of
class Magnoliopsida. Arthur Cronquist (1981) discussed Ericales under subclass Dilleniidae of
Magnoliopsida. Thorne (1983) included Ericaceae under order Ericales of superorder Theiflorae of
subclass Annonidae (Dicotyledoneae).
21.63 PRIMULALES
1. Flowers actinomorphic, bisexual, hypogynous, gamopetalous and pentamerous.
2. Stamens usually opposite the petals and epipetalous.
3. Unilocular, superior ovary with generally free-central or basal placentation.
4. The ovules are bi-integumented.

5. Seeds with straight embryo and copious endosperm.
Bentham and Hooker included Primulaceae, Plumbaginaceae and Myrsinaceae under cohort
Primulales. Engler and Diels, however, divided order Primulales into 3 families, viz. Theophrastaceae,
Myrsinaceae and Primulaceae. They discussed Plumbaginaceae under order Plumbaginales as also
did by recent workers like Takhtajan (1980), Cronquist (1981) and Dahlgren (1983).
Only Primulaceae and Plumbaginaceae have been treated in this text.
21.64 PRIMULACEAE PRIMULA OR PRIMROSE FAMILY

Gamopetalae, Heteromerae, Primulales.

Perennial herbs, only rarely shrubs; flowers pentamerous; petals fused; stamens opposite the petals;
free-central placentation.

Primula, Anagallis, Androsace, Cyclamen and Lysimachia.

Primrose family is represented by about 30 genera and 1000 species which are cosmopolitan but
chiefly distributed in northern hemisphere. Some of the larger genera with their approximately reported
species include Primula (500), Lysimachia (200), Dodecatheon (50), Dionysia (41), Anagallis (30),
Cyclamen (15), Salomus and Soldanella (11).

General Habit Mostly perennial herbs, only rarely annual herbs or shrubs; plants generally perennate
by means of rhizome (Primula) or tubers.
Leaves Mostly simple, exstipulate; sometimes variously lobed or dissected; show all the three types
of phyllotaxy i.e. alternate, opposite, or whorled; basal or cauline; often gland-dotted or farinose.
Inflorescence Flowers often borne on scapes in terminal or axillary position; solitary axillary
(Anagallis), or umbellate (Primula), or terminal racemes (Lysimachia vulgaris).
Flower Bracteate, ebracteolate, complete; actinomorphic, rarely zygomorphic (Coris); bisexual;
usually pentamerous, rarely trimerous (Pelletiera) or 5–9-merous (Trientalis); hypogynous; often
heterostyled.
Calyx Usually 5 sepals, gamosepalous (Fig. 21.75), rarely 3 to 9 sepals; valvate; persistent;
foliaceous.
370 Plant Taxonomy
stamen
gynoecium
petal
sepal
A Flower
flower
young fruit
leaf
calyx
Fruit
T.S. Ovary
stem
roots
Floral Diagram
Flowering Plant
Fig. 21.75 Anagallis arvensis L.

Corolla Usually 5 petals, gamopetalous; imbricate or quincuncial; rarely 3 to 9 petals; usually rotate
(Anagallis) to salverform (Primula); polypetalous in Pelletiera, and petals absent in Glaux; petal
lobes are reflexed sharply downward in Cyclamen (Fig. 21.76) and Dodecatheon.
calyx-lobe reflexed corolla lobe
receptacle style
stamen
ovary
L.S. Flower
A Flower
Fig. 21.76 Entire and L.S. flower of Cyclamen.
Androecium Stamens 5 or as many as corolla lobes, epipetalous; opposite the corolla lobes; some-
times 5 staminodes are also present opposite the sepals (Samolus, Soldanella); anthers dithecous,
basifixed, introrse, dehiscing longitudinally.
Gynoecium Pentacarpellary, syncarpous; ovary superior, rarely half-inferior (Samolus); unilocular,
numerous ovules; free-central placentation; style and stigma simple; heterostyly common.
Fruit and Seeds Fruit usually a 5-valved capsule or pyxis, variously dehiscent, generally dehiscing
by teeth at the tip. Seeds with small and straight embryo, in hard or fleshy endosperm.
Pollination and Dispersal Majority of the Primula species are cross-pollinated. Dispersal of seeds in
Primula is by wind but in a few species (Primula vulgaris) seeds are dispersed by ants.
General Floral Formula Br, ≈, , K(5), C (5), A5, G (5).

The family is of no specific economic importance except that of some well-known decorative
plants it contains. Some species of Primula may cause contact dermatitis. Anagallis arvensis
(Fig. 21.75) is used for curing snakebite, leprosy, gout, and even hydrophobia. Cyclamen pur-
purascence contains glucoside cyclamin which is poisonous. Some of the commonly cultivated
ornamental plants of Primulaceae are the species of Primula (primrose), Androsace (rock jas-
mine), Cyclamen (Fig. 21.76), Dodecatheon (shooting star), Douglasia, Lysimachia (loosestrile)
and Omphalogramma.
372 Plant Taxonomy

The family is divided into 4 tribes:
1. Primuleae (Ovary superior, corolla imbricate)
2. Cyclamineae (Ovary superior, corolla convolute, plants tuberous)
3. Lysimachieae (Ovary superior, corolla convolute, plants not tuberous)
4. Samoleae (Ovary half-inferior).
Because of free-central placentation, Primulaceae is related to Caryophyllaceae and also to
Theaceae to some extent. Primulaceae is also allied to Myrsinaceae and Theophrastaceae, and all
these 3 families are included in one order Primulales by many taxonomists. Bessey, Hallier and
Hutchinson opined that Primulaceae and Primulales originated from Caryophyllales. However,
Cronquist considered that Primulaceae and Primulales originated from Theales.
21.65 PLUMBAGINACEAE LEADWORT FAMILY

Gamopetalae, Heteromerae, Primulales.

Perennial herbs or shrubs, sometimes scandent or lianous; pentamerous flowers are polypetalous or
gamopetalous; 5-styled pistil; unilocular and uniovulate ovary; fruit dry, 1-seeded, surrounded by
calyx.

Plumbago, Armeria, Ceratostigma.

A family of about 19 genera and 775 species (Hickey and King, 1988), mostly of semiarid regions,
found very commonly in Mediterranean and central Asiatic regions. It shows a cosmopolitan distribu-
tion, especially along sea-shores and on salt steppes. Plumbaginaceae shows some relationship with
Primulaceae but differs from the latter in possessing solitary ovule and free styles. Most phylogenists,
however, believed that it has close affinities with caryophyllaceous taxa.

General Habit Perennial herbs or shrubs; sometimes lianous; sometimes tufted and cushion-like
(Armeria maritima).
Stem Well-branched, stout, woody.
Leaf Simple, exstipulate, entire; either forming a basal rosette of linear, punctate, glabrous, single-
veined leaves (Fig. 21.77), or arranged alternately on well-branched stems; water glands or chalk-
glands are present on the leaf surfaces.
inflorescence calyx
pedicel
C
A Floral Bud
petal
leaves
hairy
calyx
A ovary styles
(hairy)
stem
ovule
Flowering Branch ovary
F
G Gynoecium
L.S. Ovary
petals
flowers
E
stamens A Stamen
styles
ovary B
D Upper Portion of Scape

Bearing Inflorescence
L.S. Flower
Fig. 21.77 Armeria maritima (Mill.) Willd.
374 Plant Taxonomy
Inflorescence Racemose (Limonium), cymose or capitular; in Armeria maritima (Fig. 21.77) the
inflorescence is an erect scape supporting a terminal capitulum of fragrant, rose-pink or white
flowers.
Flower Bracteate; bracts scarious, sometimes forming an involucre; bisexual, actinomorphic;
pentamerous.
Calyx 5 sepals, united, persistent; sepals sometimes with smaller secondary lobes; often 5-10 ribbed
or angled or winged.
Corolla 5 petals, gamopetalous; contorted or imbricate; 5 petal lobes sometimes quite deep, appear-
ing to be polypetalous.
Androecium 5 stamens, epipetalous, opposite to corolla lobes; anthers 2-celled, dehiscing longitu-
dinally, introrse.
Gynoecium Pistil 1, ovary superior, unilocular, 5-carpelled, syncarpous, usually 5-lobed or 5-ribbed;
ovule single, pendulous, anatropous; basal placentation; styles 5, opposite to sepals, basally united,
often hairy or glandular; stigmas 5, filiform.
Fruit Dry, 1-seeded utricle, often enclosed within persistent calyx.
Seed Embryo straight, with floury or crystalline-granular endosperm.
General Floral Formula Br, ≈, , K(5), C (5) or 5, A5, G (5).

Except of the ornamental value of some plants, the family Plumbaginaceae is of little importance.
Ornamental plants include some species of Armeria (Thrift or Sea pink), Limonium (Satice),
Plumbago, Ceratostigma and Acantholimon. Dried floral branches of Limonium are used in floral
decoration while Plumbago indica and P. auriculata are cultivated as semi-climbing subshrubs in
greenhouses. Some species of Plumbago and Limonium are used as medicine. Plumbago euro-
paea and P. scandens are used to treat dental ailments. Extracts from the leaves and roots of P.
zeylanica are used to treat some skin diseases.

Plumbaginaceae shows some relationship with Primulaceae due to a common floral plan with
antipetalous stamens, unilocular ovary and bitegmic ovule but is distinguished from Primulaceae
by its solitary ovule and free styles. Hutchinson considered it as one of the two families of
Primulales. Lawrence believed that there is evidence that the Plumbaginales (having a single family
Plumbaginaceae) “may have evolved from stocks ancestral to the Primulales, and most phylogenists
have been of the opinion that they have close affinities with the caryophyllaceous taxa”. However, both
orders (Primulales and Plumbaginales) have been “derived from Centrospermae or their ancestors”.
21.66 EBENALES
1. Flowers gamopetalous.
2. Stamens usually in 2–3 whorls, epipetalous.
3. Ovary superior.
4. Carpels more than two, usually 4 or 5, syncarpous, superior.
5. Ovary usually with as much locules as number of carpels, each locule is generally
uniovulate.
6. Placentation axile.
Bentham and Hooker included 3 families in Ebenales, namely Sapotaceae, Ebenaceae and
Styraceae. Engler and Diels, however, treated the order Ebenales as composed of 2 suborders
and 7 families; suborder Sapotineae (Sapotaceae, Hoplestigmataceae) and suborder Diospyrineae
(Ebenaceae, Diclidantheraceae, Symplocaceae, Styracaceae, Lissocarpaceae).
Only Sapotaceae is discussed here in some details.
21.67 SAPOTACEAE SAPOTA FAMILY

Gamopetalae, Heteromerae, Ebenales.

Plants with milky latex; leaves exstipulate; stamens in 2–3 whorls; ovary superior; axile placenta-
tion; fruit berry.

Achras, Madhuca, Manilkara, Mimusops.

Family includes about 40 genera and over 800 species, with their main distribution in tropical
regions of the world. Only 12 genera and about 50 species have been reported from India, mainly
from eastern and southern regions. Because of Sapodilla (Achras sapota), the family is also called
“Sapodilla family”.

General Habit Mostly trees or shrubs, with milky sap present in the laticiferous ducts or sacs in
their vegetative parts.
Leaves Simple, alternate, petiolate; exstipulate but rarely stipulate (Madhuca); lanceolate, obcordate,
or elliptical; entire; tomentose (Madhuca); unicostate reticulate.
Inflorescence Usually cymose clusters; sometimes solitary axillary (Chrysophyllum) or solitary
terminal; rarely cauliflorous (Dichopsis).
Flower Bracteate (Madhuca) or ebracteate (Achras), pedicellate, bisexual, actinomorphic,
hypogynous.
Calyx 4 to 8 sepals, usually gamosepalous, sometimes free; imbricate or valvate; 6 sepals of
Palaquium, Mimusops and Achras are arranged in 2 series of 3 each; in Madhuca indica 4 sepals
are arranged in 2 whorls of 2 each.
376 Plant Taxonomy
leaf
T.S. Ovary
leaf
fruit
stamens
corolla
stem
Flowering Branch Expanded Corolla

Floral Diagram
Fig. 21.78 Madhuca indica Gmel.
Corolla Petals usually as many as sepals, gamopetalous, imbricate; sometimes petals contain dorsal
or lateral appendages; sometimes petals are twice as many as sepals (Madhuca indica, Fig. 21.78);
only 6 petals in Achras sapota.
Androecium Stamens many, usually in 2 or 3 whorls of 4–5 each; often the stamens of innermost
whorl are fertile while that of all others are reduced to staminodes (Mimusops); epipetalous, poly-
androus; filaments short; anthers dithecous, basifixed or dorsifixed, extrorse, with their connective
often prolonged beyond anther lobes, dehiscing longitudinally.
Gynoecium Carpels varying from 1–14 but usually 4 or 5, syncarpous, superior; usually with as
many locules as the number of carpels, each locule uniovulate, ovule anatropous, axile placentation;
style 1, often with apically lobed stigma.
Fruit and Seeds Fruit often berry; seeds usually endospermic, with straight embryo.
General Floral Formula Br, ≈, , K(4–8), C (4–8) or Aμ, G (1–14).

• Chewing gum: Chicle for chewing gum is obtained from the coagulated resinuous latex
obtained from the bark of Achras sapota (Sapodilla plum)
• Gutta percha: Gutta percha, used for insulation and other similar purposes, is obtained
from several genera, such as Palaquium (P. gutta), Payena, Mimusops, etc.
• Edible fruits: Notable edible fruits of Sapotaceae include Achras sapota (Sapodilla plum or
Chiku), Manilkara hexandra (Khirni), M. kauki (Kauki), Chrysophyllum cainito (star apple),
Pouteria campechina (eggfruit), Calocarpum sapota (Sapote plum), etc. Dried corolla of
Madhuca indica is eaten by man.
• Timber: Valuable timber is obtained from Manilkara hexandra, Sideroxylon, Madhuca
indica (Mahua), Xantolis tomentosa, etc.
• Medicinal value: (i) Chicle gum, obtained from Achras sapota, is used in dental surgery,
(ii) Seeds and dried fruits of Mimusops elengi (Maulsari) are highly effective in curing piles,
(iii) Oil from seeds of Madhuca indica is effective in skin diseases and rheumatism.
• Oil: Oil, obtained from seeds of Madhuca indica and Diploknema butgracea, is used in
the manufacture of soaps and candles.

Sapotaceae is usually divided into 2 tribes, viz. Palaquieae (petal lobes without appendage) and
Mimusopeae (petal lobes with appendage).
Sapotaceae is allied closely to Ebenaceae, and both these families belong to order Ebenales.
However, Hallier placed Sapotaceae under a monotypic order Sapotales, while Ebenaceae under
Santalales. Majority of the phylogenists consider that both Sapotales and Santalales originated from
the same ancestral stock, i.e. Linaceae.
21.68 GENTIANALES
1. Leaves opposite, simple or pinnately compound and exstipulate.
2. The flowers are bisexual, actinomorphic and hypogynous.
3. Corolla gamopetalous and corolla lobes usually convolute.
4. Androecium of one series; stamens adnate to or near corolla base, epipetalous and
alternipetalous.
5. Gynoecium bicarpellary and ovary superior.
Bentham and Hooker included 6 families under Gentianales viz. Oleaceae, Apocynaceae,
Asclepiadaceae, Salvadoraceae, Loganiaceae and Gentianaceae. Engler and Diels used the name
“Contortae” for Gentianales and divided it into 2 suborders containing 6 families: (i) suborder
Oleineae (Oleaceae) and, (ii) suborder Gentianineae (Desfontaineaceae, Loganiaceae, Gentianaceae,
Apocynaceae, Asclepiadaceae).
Oleaceae, Asclepiadaceae, Apocynaceae and Loganiaceae have been discussed in this text.
21.69 OLEACEAE OLIVE FAMILY

Gamopetalae, Bicarpellatae, Gentianales.
378 Plant Taxonomy

Trees or shrubs; leaves opposite; flowers bi- to tetramerous; gamosepalous and gamopetalous;
stamens 2; ovary bilocular.

Nyctanthes1, Jasminum, Fraxinus, Myxopyrum, Osmanthus and Olea.

A family of approximately 29 genera and 600 species, Oleaceae are cosmopolitan but occur in great
diversity in temperate and tropical Asia. Some of the largely represented genera of olive family
and their approximately reported species are Jasminum (300), Chionanthus (125), Linociera (80),
Fraxinus (70), Ligustrum (40), Syringa (30) and Olea (20).

General Habit Shrubs (Syringa) or trees (Olea, Nyctanthes), occasionally vines (Myxopyrum
smilacifolium).
Leaves Simple (Nyctanthes) or pinnately compound (Fraxinus); usually opposite, rarely alternate
(some species of Jasminum); often entire, exstipulate; unicostate reticulate venation.
Inflorescence Axillary or terminal racemose, paniculate or thyrsiform.
Flower Bracteate (Olea) or ebracteate (Jasminum), pedicellate, usually bisexual but unisexual in
some species of Olea and Fraxinus (Fig. 21.79G); actinomorphic, bi- to hexamerous but usually
tetramerous, hypogynous.
Calyx 4 sepals, gamosepalous, valvate; rarely sepals absent (Fraxinus excelsior, Fig. 21.71F); some-
times 4–15 toothed; persistent.
Corolla Usually 4 united petals, valvate or imbricate; sometimes 4–9 lobed (some species of
Jasminum); rarely petals absent (Fraxinus micrantha); sometimes deeply lobed or divided and seem-
ingly polypetalous.
Androecium Usually 2 stamens, epipetalous, alternate with the carpels; anthers dithecous, the cells
usually back to back; often apiculate because of the extension of the connective; dehiscing longitu-
dinally; stamens are 4 in Hesperelaea and Tessarandra.
Gynoecium Usually bicarpellary, syncarpous; ovary superior, bilocular; with usually 2 anatropous
ovules in each locule; axile placentation; style simple, stigma capitate or bilobed.
Fruit and Seeds Fruit drupe (Olea), berry (Ligustrum), samara (Fraxinus, Fig. 21.79H), or capsule
(Forsythia) with 1–4 seeds. Seeds with straight embryo and little or no endosperm.
General Floral Formula Br or Ebr, ≈, , K(4), C (4), A2, G (2).
1
Some workers treat Nyctanthes as a member of sub-family Nyctanthoideae of family Verbenaceae.
stamen ovary
G
Male Flower
E
F
Flowering Branch Bisexual Flower
stigma
inflorescence
stamen H
Fruit
petal (samara)
style
ovary
C
B L.S. Flower
A Flower
leaf
stem
A Floral Diagram
Flowering Branch
Fig. 21.79 A–D: Nyctanthes arbortristis, and E–H: Fraxinus excelsior.

380 Plant Taxonomy

Olive family is of considerable economic importance as a source of olives, olive oil, ash lumber,
dyes and several ornamental plants.
• Notable ornamentals are various species of Jasminum (Jasmine), Syringa (lilac), Forsythia
(golden bells), Ligustrum (privet), Osmanthus (fragrant olive), Phillyrea, Chiananthus (fringe
tree) and Nyctanthes.
• Olive (Olea europaea) has been a source of food since ancient times, and the oil extracted
from its fruit is of high value.
• Flowers of several species of Jasminum are the source of an oil used in perfumery.
• Valuable timber is obtained from various species of Fraxinus (ash lumber), Notalaea and
Olea.
• An orange dye, used for colouring silk and cotton, is obtained from the flowers of various
species of Nyctanthes. Leaves of N. arbortristis (Har Singar, Fig. 21.79A–D) are useful in
fever and rheumatism.

Family Oleaceae is divided into 2 subfamilies, i.e. Jasminoideae (ovules generally erect) and
Oleoideae (ovules pendulous). To some extent, Oleaceae is related to Loganiaceae. In certain
characters it is also allied to Apocynaceae. According to Hutchinson, Oleaceae shows affinities
to Melastomaceae and Rubiaceae. However, Cronquist placed Oleaceae under Scrophulariales and
believed it to be allied to Buddleiaceae. According to Takhtajan, Oleaceae belongs to a separate
order Oleales and is closely related to some families of order Celastrales. Takhtajan regards that both
Oleales and Celastrales have a common origin from Saxifragales.
21.70 ASCLEPIADACEAE MILKWEED FAMILY


Herbs or vines with milky sap; leaves exstipulate and opposite or whorled; flowers pentamerous
with bicarpellate gynoecium; corona, pollinia, translators and corpuscula are present; fruit follicle;
silky hairs cover the seeds.

Asclepias, Calotropis, Cryptostegia, Leptadenia, Tylophora.

A family of about 250 genera and 2000 species (Jones and Luchsinger, 1987), distributed mainly in
tropical and subtropical regions and a few also in temperate regions. About 50 genera and over 250
species of Asclepiadaceae have been reported from India. Some of the larger genera along with their
number of approximately reported species are Hoya (200, wax plant), Ceropegia (160), Asclepias
(150, milkweed), Oxypetalum (150), Matelea (130, milkweed vine), Dischidia (80), Stapelia (75,
carrion flower) and Periploca (10).

General Habit Erect or twining perennial herbs (Asclepias), shrubs (Calotropis procera), vines
(Leptadenia), or rarely small trees (Calotropis gigantea); usually with milky sap; often fleshy or
cactus-like (Stapelia), sometimes epiphytic (Dischidia). Rootstock fleshy, tuberous, woody, or some-
times absent.
Leaves Simple; usually opposite decussate (Calotropis procera, Fig. 21.82), sometimes alternate
or in whorls; reduced or even absent in succulent species; usually exstipulate, or stipules minute if
present; margins generally entire; often have a waxy coating.
Inflorescence Mostly cymose (dichasial to multichasial cyme), but also racemose or umbellate
(Stapelia).
Flower Bracteate, often bracteolate, complete; actinomorphic, rarely zygomorphic (Ceropegia);
hermaphrodite, pentamerous, hypogynous, cyclic.
Calyx 5 free or basally united sepals forming a short tube; valvate, imbricate, or quincuncial.
Corolla 5 petals, gamopetalous; contorted, imbricate or valvate; corolla tube often short and
with lobes; corolla tube is zygomorphic in Ceropegia, funnel form in Cryptostegia and campanu-
late in Gymnema; often the corolla tube is with a corona, made up of a ring of scales, hairs or
processes.
Androecium 5 stamens, epipetalous, rarely distinct, pollen in waxy pollinia.
Typically, the filaments of stamens are connate into a short tube and their anthers are usually
adherent or adnate to the stigmatic area of the gynoecium to form a complex structure called gynos-
tegium or gynandrium (Fig. 21.80A, B). Anthers are 2-celled and their pollens often united within
each anther sac into a structure called pollinium. Two pollinia, one each from the adjacent anthers,
are united together by two arms (retinaculae and translators) attached to a central gland called
corpusculum (Fig. 21.80C).
A nectariferous corona, consisting of 5 hoods, may be present. These hoods are sometimes mis-
taken for petals. A beak or hood may be associated with each hood of corona (Fig. 21.80B).
Gynoecium Bicarpellary; the carpels are free below but united by their apices into the single 5-lobed
stigma; ovary superior; ovary of each carpel is unilocular; numerous ovules in each locule, marginal
placentation (Fig. 21.81A); styles 2; stigma 1 and 5-lobed.
Fruit and Seeds Fruit is a pair of follicles (Fig. 21.81B), of which often one aborts. Seeds numer-
ous, with tufts of silky hairs called coma (Fig. 21.81C); embryo straight, large, nearly as long as the
seed, with small and thin endosperm.
Pollination It is entomophilous, and butterflies are the chief pollinating agents which visit the
brightly coloured flowers in search of nectar. When a butterfly alights on a flower in search of
nectar, its legs tend to slip down the slits in the column and hook onto the band connecting a pair
of pollinia. As the butterfly flies away, this connecting band twists sharply. This brings the pollinia
382 Plant Taxonomy
cucullus corona-horn cucullus corona-horn
gynostegium
stigma
pollinium
stigmatic
groove
corpusculum
translator
column column
style style pollinium retinaculum

pollen
ovary B C mass
A
Translator and A
Pair of Pollinia
corolla lobes
Corolla, Column
L.S. Flower (central part) & Corona
Fig. 21.80 A–C Floral details of Asclepias curassavica.
ovules
hairs
seeds
A B C
Mature Fruit A Seed
L.S. Carpel
(lower part)
Fig. 21.81 A–C Carpel, fruit and seeds of Asclepias curassavica.
close together. When the butterfly arrives at another flower, the pollinia are in such a position that
they lodge in a slit in the column of the flower and break off, and thus distributing the pollen grains
over the stigma.
General Floral Formula Br, ≈, , K(5) or 5, C (5), A(5), G (2).

• Ornamental plants of this family grown in glasshouses include several species of Asclepias,
Caralluma, Ceropegia, Cryptostegia, Hoya, Huernia, Oxypetalum, Periploca, Stapelia and
Stephanotis.
corpusculum
stigmatic disc
caudicle
pollinium
pollinium
flower bud style
A Stamen ovary
fruit
leaf
Gynoecium
stem
hairs Flowering Branch
Seed
Fruit
pollinium stigmatic disc
anther
cucullus
style
ovary
ovule sepal
pedicel
Floral Diagram
L.S. Flower
Fig. 21.82 Calotropis procera R. Br.
• Fibre, used for cordage, twines, fishing nets, etc., is obtained from Asclepias curassavica
(blood-flower), Calotropis gigantea, Cosmostigma racemosa, Leptadenia pyrotechnica,
Marsdenia roylei, M. tinctoria and Tylophora tenuis.
• Indigo-like dye is obtained from Marsdenia tinctoria.
• Cryptostegia grandiflora has been cultivated commercially as a source of rubber.
• Latex obtained from species of Matelea is used as an arrow poison.
• Certain plants of Asclepiadaceae are of medicinal importance. These include, (i) Calotropis
procera (roots are used against cough), (ii) Gymnema sylvestre (used as a laxative
384 Plant Taxonomy
and diuretic), (iii) Hemidesmus indicus (roots are used in urinary and skin diseases),
(iv) Holostemma rheedianum (roots are used in cough and cold), (v) Pergularia daemia
(used as an emetic and expectorant), (vi) Sarcostemma acidum (infusion of roots is used as
an antidote for snakebite), and (vii) Tylophora indica (roots are used in curing bronchitis,
whooping cough and asthama).

On the basis of pollen characters, Asclepiadaceae is divided into two subfamilies, viz. (i) Periplocoideae
(pollen grains in tetrads; filaments free), and (ii) Cynanchoideae (pollen grains united in pollinia;
filaments united in a tube).
Asclepiadaceae is most closely allied to Apocynaceae, and in majority of the systems of clas-
sification both these families are placed side by side. However, the characters such as presence of
translators and presence of gynostegium are absent in Apocynaceae.

Calotropis procera R. Br. (Madar or Aak, Fig. 21.82)
Habit: A herbaceous or semishrubby weed with its upper portions covered with wooly hairs; contains
milky latex. Leaf: Simple, opposite decussate, exstipulate, sessile or subsessile, ovate to oblong, entire,
acute, unicostate reticulate. Inflorescence: Dichasial cyme. Flower: Bracteate, bracteolate, pedicel-
late, hermaphrodite, actinomorphic, pentamerous, hypogynous, cyclic. Calyx: 5 sepals, distinct,
quincuncial. Corolla: 5 petals, fused, twisted, coloured. Androecium: 5 stamens, united with stigma
to form gynostegium; each stamen is represented by two pollinia with their caudicles or retinaculae;
pollinia of the adjacent anthers are joined by their retinaculae to corpusculum in a groove, to form
a unit called translator; at the back of each stamen is present a coronary outgrowth. Gynoecium:
Bicarpellary, superior; ovaries free but the upper portions of styles and stigma fused; each ovary is
unilocular, many ovules in the locule, marginal placentation; styles 2; stigmatic head pentagonal.
Fruit: Follicle. Floral Formula: Br, Brl, ≈, , K5, C (5), A(5), G2.
21.71 APOCYNACEAE DOGBANE FAMILY


Plants with milky sap; leaves opposite or whorled, exstipulate; flowers pentamerous; stamens usu-
ally epipetalous; corona, pollinia and corpusculum of Asclepiadaceae absent; fruit follicle and seeds
with hairs.

Apocynum, Catharanthus, Nerium, Plumeria, Tabernaemontana, Thevetia, Rauvolfia, Vinca.

A family of about 200 genera and 2000 species (Jones and Luchsinger, 1987) of mostly tropical and
subtropical distribution, but a few species distributed also in temperate regions. About 30 genera and
over 60 species of Apocynaceae have been reported from India. Some of the larger genera along
with their approximately reported species in bracket are Rauvolfia (100), Tabernaemontana (100),
Parsonsia (100), Aspidosperma (80), Prestonia (65), Strophanthus (60), Apocynum (7), Plumeria
(6), Vinca (5) and Catharanthus (5).

General Habit Trees (Alstonia, Holarrhena), shrubs (Thevetia, Nerium), herbs (Catharanthus),
woody twiners (Allamanda), or even succulent (Adenium); usually twining shrubs with milky sap
in laticiferous vessels.
Leaves Simple, exstipulate; opposite decussate or sometimes alternate (Thevetia) or in whorls of 3;
entire; rarely stipulate (Tabernaemontana).
Inflorescence A panicle, cyme, or raceme, or flowers solitary (Catharanthus); usually dichasial
cyme.
Flower Bracteate, bracteolate, complete, hermaphrodite, actinomorphic, pentamerous; hypogynous
but rarely perigynous or epigynous (Plumeria).
Calyx 5 sepals, usually gamosepalous, sometimes free (Thevetia, Catharanthus) or deeply lobed;
valvate (Catharanthus), quincuncial (Thevetia); odd sepal posterior; rarely only 4 sepals; sometimes
glandular appendages (Squamellae) present on the adaxial side.
Corolla 5 petals, united into a tube, usually salver-shaped or funnel-shaped; twisted or rarely val-
vate; rarely only 4 petals; often hairy within or contain some corona-like outgrowths at the mouth
of corolla tube.
Androecium 5 stamens, epipetalous; alternate with the corolla lobes; filament short; attached at
the mouth or deeper in the corolla tube; anthers are dithecous, basifixed, often sagittate; introrse;
dehiscence longitudinal; anthers sometimes basally awned; sometimes bear hairy appendage (Nerium)
over the lobes (Fig. 21.83).
Gynoecium Usually bicarpellary, carpels apically united; superior or rarely half-inferior (Plumeria);
1- or 2-locular, with 2 to many anatropous and pendulous ovules in each locule; marginal placen-
tation; style one and simple; stigma is characteristically thickened, massive and bilobed; receptive
surface of stigma is situated below; rarely the carpels are 3–5.
A nectariferous disc is often present around or at the base of the gynoecium in several genera,
e.g. Thevetia, Catharanthus, Allamanda and Rauvolfia.
Fruit and Seeds Fruit usually consisting of 2 follicles, or a berry (Landolphia), drupe (Cerbera),
capsule (Allamanda), or 2 indehiscent mericarps. Seeds often flat with straight embryo and a crown
of hairs.
Pollination and Dispersal Pollination is entomophilous. Insects are attracted mainly because of often
large and conspicuous petals and the presence of nectar. Because the receptive stigmatic surface is
located on the underside of the stigma, the cross-pollination is the rule of the family. Self-pollination
is absent.
386 Plant Taxonomy
ovary wall
locule
appendage
ovule
anther lobe
T.S. Ovary
filament
Stamen
flower
inflorescence
Floral Diagram
appendage
stigma stigma
petal
style stamen
style
leat
ovary ovary
sepal
pedicel pedicel
stem
Gynoecium L.S. Flower
Flowering Branch
Fig. 21.83 Nerium indicum Mill.
Dispersal is affected mainly by wind because of winged seeds and presence of crown of hairs.
General Floral Formula Br, Brl, ≈, , K(5), C (5), A5, G (2).

The family is of importance for its several ornamentals and for drugs.
• Ornamental plants1 include Allamanda neriifolia (golden yellow flowers), Alstonia scholaris
(pink flowers), Beaumontia grandiflora (large white flowers), Carrisa carandas (white-pink
1
Plants arranged alphabetically.
flowers), Catharanthus roseus syn. Vinca rosea (Sadabahar, rosy-purple flowers), Mandevilla
taxa (pink flowers), Nerium indicum syn. N. odorum (Kaner, white-pinky flowers), Plumeria
(white, red, pink or purple flowers), Tabernaemontana divaricata (Chandni, white flowers),
Thevetia peruviana (Peeli Kaner, yellow flowers), and Vallaris solanacea (Dudhi Bel).
• Plants of medicinal value include (i) Rauvolfia serpentina (Sarpgandha) used in the treatment
of hypertension and mental disorders; drug is obtained from the dried roots; the alkaloid
reserpine of the plant can lower the blood pressure and tranquilize mental patients suffering
from schizophrenia, (ii) Thevetia peruviana (seeds are used in rheumatism), (iii) Vallaris
solanacea (latex is useful in toothache and inflated gums), (iv) Holarrhena antidysentrica
(root and bark is used in curing dysentery), (v) Alstonia scholaris (bark is used in malaria and
dysentry), (vi) Cerbera manghas (latex is used as an emetic and purgative), (vii) Strophanthus
hispidus (seeds yield the drug strophanthin), and (viii) Wrightia tomentosa (bark and roots
are used as antidote to snakebite).
• Rubber is obtained from the coagulated latex of several species of Carpodinus, Funtumia,
Hancornia, Landolphia and Rhynchodia.
• Poisonous plants of the family belong to Acokanthera, Nerium, Strophanthus and Thevetia.
If one leaf of Nerium oleander or Thevetia peruviana is ingested, it may be fatal to an
adult human. Poisoned arrows are made from the bark extract of Acokanthera abyssinica
and seeds of several species of Strophanthus.
• Fruits of Carissa carandas (Karonda) are edible and used in pickles.
• An indigo-like dye is obtained from the seeds of Wrightia tinctoria.

The family is closely allied to Asclepiadaceae and both are included in the same order almost in all
systems of classification. However, Apocynaceae lack the pollinia, corona and translator-corpusculum
of Asclepiadaceae.

Catharanthus roseus (L.) Don. syn. Vinca rosea Reichb. (Periwrinkle or Sadabahar; Fig. 21.84)
Habit: Erect, everblooming ornamental plant with milky latex. Leaf: Simple, opposite decus-
sate, exstipulate, subsessile or petiolate, elliptic-obovate, entire, mucronate, unicostate reticulate.
Inflorescence: Dichasial cyme or solitary axillary. Flower: Ebracteate, pedicellate, complete, her-
maphrodite, actinomorphic, pentamerous, hypogynous, rosy-purple or pink. Calyx: 5 sepals, polyse-
palous, valvate. Corolla: 5 petals, gamopetalous, twisted, hypocrateriform. Androecium: 5 stamens,
epipetalous, inserted at the mouth of corolla tube, filaments short; anthers sagittate, dithecous, bas-
ifixed or dorsifixed, introrse. Gynoecium: Bicarpellary, syncarpous; ovaries free and superior; many
ovules, marginal placentation; two hypogynous nectaries are present, one on anterior and another on
posterior side of the ovary. Fruit: Follicle.
Floral Formula: Ebr, ≈, , K5, C (5), A5, G (2).
388 Plant Taxonomy
anther
lobe
connective
filament
Stamen
petal
stigmatic Floral Diagram

stamen head
flower
style
bud
nectary
ovary
sepal fruit
ovule
thalamus
pedicel Gynoecium
L.S. Flower
ovary wall
nectary
locule
ovule
leaf
placenta
stem
T.S. Ovary
Flowering Branch
Fig. 21.84 Catharanthus roseus (L.) Don. syn. Vinca rosea Reichb.
21.72 LOGANIACEAE LOGANIA FAMILY

Gamopetalae, Bicarpellatae, Gentianales

Leaves opposite, stipulate; ovary superior, bilocular; axile placentation; most members show the
presence of internal phloem.

Logania, Strychnos, Buddleia, Fagraea.

A family of over 32 genera and about 800 species, Loganiaceae are distributed mainly in tropical
regions. A few members are also found in warm temperate regions. Some of the larger genera along
with their approximately reported species are Strychnos (200), Buddleia (150), Geniostoma (60),
Nuxia (40), Spigelia (35) and Logania (25).

General Habit Herbs, shrubs or trees; several members are climbers.
Leaves Usually simple, opposite; stipulate, the stipules often much reduced; in a few species of
Buddleia, the leaves are alternate or even whorled; lanceolate and finely toothed in Buddleia davidii
(Fig. 21.85).
Inflorescence Usually cymose.
Flower Bracteate, bracteolate, bisexual, actinomorphic; mostly tetra- to pentamerous.
Calyx Sepals 4 or 5, gamosepalous, imbricate.
Corolla Petals 4 or 5, gamopetalous; rarely 10-lobed corolla; showing various types of aestivation;
a crown of hairs often present at the mouth of corolla tube; corolla lobes have irregular margins as
in Buddleia davidii (Fig. 21.85).
Androecium Stamens 4 or 5, generally as many as corolla lobes, epipetalous, alternipetalous; rarely
stamens twice as many as petals, sometimes only 1 stamen (Usteria); anther lobes dithecous; dehisc-
ing longitudinally.
Gynoecium Usually bicarpellary, syncarpous; superior or rarely half-inferior (e.g. Mitreola) ovary,
bilocular; usually numerous anatropous or amphitropous ovules in each locule on axile placentation;
the ovary is unilocular in Strychnos or incompletely bilocular in Fagraea; style 1, rarely 2 (e.g.
Cynoctonum); stigmas 1–2.
Fruit Usually a septicidal capsule, rarely drupe or berry.
Seeds Endospermic with small and straight embryo; sometimes winged .
General Floral Formula Br, Brl, ≈, , K(4 – 5), C (4–5) or A4–5 , G (2).
390 Plant Taxonomy
petals
flowers
corolla
tube
leaf
stem
A
sepals
Inflorescence
bract
B
A Single Flower
ovules
locule
F
D T.S. Ovary
G A Young Anther
A Fruit stigma
corolla
tube corolla
tube style
stamens
sepal
hairs ovary
(superior)
filaments
adnate to
tube
E
C
L.S. Flower
Corolla Tube Opened Out to (Lower Portion)
Show Stamens
Fig. 21.85 Buddleia davidii Franch.

• Strychnos nux-vomica, the source of alkaloids like strychnine and brucine, is well-known
as a drug plant. The famous drug nux-vomica, obtained from its seeds, is used as a tonic,
stimulant and in the treatment of nervous disorders and paralysis, but used always in small
doses because it is a virulent poison.
• The drug, curare, used in shock therapy, as a muscle relaxant and also in surgical operations
and tetanus, is obtained from the bark of Strychnos toxifera.
• Fruits of Strychnos spinosa are edible.
• Several species of Buddleia (butterfly bush), Gelsemium (yellow jessamine), Spigelia (Indian
pink) and Logania are grown for decorative purposes.

Bentham and Hooker included 6 families (Oleaceae, Salvadoraceae, Asclepiadaceae, Loganiaceae,
Apocynaceae and Gentianaceae) under order Gentianales. Hutchinson, however, discussed Loganiaceae
and Oleaceae under order Loganiales of division Lignosae. Most taxonomists included Loganiaceae
in the Contortae, except Hallier, who placed it in Tubiflorae. Bessey suggested that Gentianales
arose from Geraniales, which separated into two phyletic lines, one giving rise to Loganiales and
the other to the Polemoniales.
21.73 POLEMONIALES
1. It includes mostly herbs and climbers; only some are shrubs and trees.
2. Leaves are usually simple and alternate.
3. Flowers actinomorphic, only rarely zygomorphic; bisexual, hypogynous, pentamerous.
4. Corolla gamopetalous, imbricate.
5. Stamens are epipetalous and alternipetalous; usually adnate to the mostly elongated corolla
tube.
6. Gynoecium with 2–5 carpels; syncarpous, superior.
Bentham and Hooker included five families in Polemoniales. These are Boraginaceae, Polemoniaceae,
Hydrophyllaceae, Convolvulaceae and Solanaceae. Engler and Diels, however, included all these
five families under order Tubiflorae, to which they divided into 8 suborders and 23 families. They
included families Convolvulaceae and Polemoniaceae under suborder Convolvulineae, families
Hydrophyllaceae and Boraginaceae under suborder Boragineae, and to family Solanaceae under
suborder Solanineae. Several families (e.g. Verbenaceae, Labiatae, Scrophulariaceae, Bignoniaceae,
Pedaliaceae and Acanthaceae), discussed elsewhere in this text, have also been included by Engler
and Diels under order Tubiflorae.
Only Boraginaceae, Convolvulaceae, Solanaceae and Polemoniaceae have been discussed in this
text.
21.74 BORAGINACEAE BORAGE FAMILY

Gamopetalae, Bicarpellatae, Polemoniales.
392 Plant Taxonomy

Bristly or hispid herbs; leaves alternate; inflorescence helicoid cyme; flowers actinomorphic,
pentamerous; ovary 4-lobed; style gynobasic; fruit of 4-nutlels.

Heliotropium, Cynoglossum, Trichodesma, Ehretia and Cordia.

A family of approximately 100 genera and over 2,000 species, distributed in both tropical and
temperate regions but mainly in the Mediterranean region. About 37 genera and over 150 species
of Boraginaceae have been reported from India, mainly from the temperate regions and alpine
Himalayas. Some of the larger genera along with their number of approximately reported species
are Heliotropium (250), Tournefortia (150), Cryptantha (100), Plagiobothrys (l00), Eritrichium
(65), Lithospermum (60), Lappula (55), Cynoglossum (50–60), Mertensia (50), Myosotis (50) and
Amsinckia (50).
Myosotis, commonly named as “forget-me-not”, perhaps derived its name from the “last words of
a gallant gentleman who drowned while crossing a stream to collect the bouquet of these sky-blue
flowers for his girl friend” (Jones and Luchsinger, 1987).

General Habit Usually bristly or hispid, hairy, herbaceous (Heliotropium) plants with fleshy roots
or rhizomes; sometimes shrubs (some species of Tournefortia), trees (Ehretia, Cordia) or lianas
(some species of Cordia).
Leaves Usually simple, alternate, exstipulate; margin usually entire; often covered by rough hairs;
rarely opposite (Trichodesma); usually narrow and sessile (Heliotropium).
Inflorescence Usually a scorpioid or helicoid cyme or a coiled “cincinnus’ uncoiling as flowers
open so that newly opened flowers face the same direction; sometimes panicle or corymb (Ehretia,
Cordia).
Flower Bracteate, bisexual, actinomorphic (rarely zygomorphic, e.g. Lycopsis, Echium); pentamer-
ous, hypogynous.
Calyx 5 sepals, distinct (Lithospermum) or basally connate to form a short or long tube; quincuncial
(Heliotropium) or imbricate, or rarely valvate:
Corolla 5 petals, gamopetalous, often lobed; imbricate (Heliotropium) or twisted; tubular, infundibu-
liform, campanulate or rotate (Trichodesma); often with projecting scales (Heliotropium) in throat
in the alternate position with the stamens (Fig. 21.86).
Androecium 5 stamens, epipetalous, alternate with petals; anthers dithecous; sagittate or conical;
basifixed, introrse, longitudinally dehiscent; nectariferous disc sometimes present at the base of
filaments.
Gynoecium Bicarpellary, syncarpous; ovary superior, bilocular, with 2 ovules in each locule; often
becoming 4-locular and deeply 4-lobed by the growth of false septum, with each locule containing
petals
petal
stamen
sepal
stigma
sepal Flower
style
ovary
L.S. Flower
Floral Diagram
Flowering Plant
Fig. 21.86 Heliotropium strigosum Willd.
one ovule; ovules anatropous, erect, ascending or horizontal; placentation axile, sometimes appearing
basal; style usually gynobasic; stigma typically one.
Fruit and Seeds Fruit usually of 4 achenes (nutlets), or a drupe, or a nut. Seeds with erect or curved
embryo and usually non-endospermic.
Pollination It is entomophilous. Insects are attracted because of the presence of nectar, colour of
the flowers and floral aggregation on the upper part of the inflorescence branches.
General Floral Formula Br, ≈, , K(5) or 5, C (5), A5, G (2).

Except that of some ornamental and medicinal value, the Borage family is not of much economic
importance.
• Ornamental plants1 of the family include species of Anchusa (Alkanet), Borago (Borage),
Cordia (Geiger tree), Cynoglossum (Hound’s-tongue), Echium (Viper’s Bugloss), Heliotropium
1
394 Plant Taxonomy
(Heliotrope), Mertensia (Blue bells), Myosotis (Forget-me-not), Omphalodes (Creeping

forget-me-not), Pulmonaria (Lungwort) and Symphytum (Comfrey).
• Plants1 of medicinal value include (i) Cordia myxa (bark and fruits are used in coughs),
(ii) Ehretia macrophylla (leaves are used in cough and stomach trouble), (iii) Heliotropium
indicum (leaf-decoction is used in fever, and roots in cough), (v) Lithospermum officinale
(seeds are used in urinary bladder diseases), and (v) Onosma bracteatum (leaves and flowers
are used as tonic and diuretic).
• Edible plants of the family are (i) Cordia dichotoma (Lassora), of which the unripe fruits
are used as vegetable, and (ii) Cordia gharaf (Gondhi), of which fruits are edible.
• Ratanjot, a red dye, is obtained from the roots of Onosma hispida.
• Wood of various species of Ehretia is used in making match boxes, brush backs, tea boxes,
and other similar articles.

Willis (1973) divided Boraginaceae into 2 sub-families viz. Heliotropioideae (style terminal; fruit
drupe) and Boraginoideae (style gynobasic; fruit achenes). Boraginoideae is further divided into 5
tribes, i.e. Cynoglosseae, Eritricheae, Boragineae, Lithospermeae and Echieae.
Takhtajan (1969) placed Boraginaceae under Polemoniales, while Cronquist (1981) placed it under
order Lamiales, and Thorne (1983) under Solanales. Hutchinson placed Boraginaceae under a mono-
typic order Boraginales. The family is allied closely to Polemoniaceae, Hydrophyllaceae, Solanaceae
and Convolvulaceae. According to Hallier, Boraginaceae is derived directly from Annonaceae.
21.75 CONVOLVULACEAE MORNING GLORY FAMILY


Usually climbing or twining vines with milky sap; leaves alternate, exstipulate; flowers axillary
and pentamerous; corolla tubular, infundibuliform or salverform; stamens 5, epipetalous; gynoecium
bicarpellary.

Argyreia, Convolvulus, Cuscuta, Evolvulus, Ipomoea.

A family of about 55 genera and 1650 species, distributed in both tropical and temperate regions of
the world. About 20 genera and over 150 species have been reported from India. Some of the largely
represented genera with their number of approximately reported species are Ipomoea (500, Morning
1
glory), Convolvulus (250, blindweed, or Wild Morning glory), Cuscuta (170, Dodder), Jacquemontia
(120), Evolvulus (100) and Calystegia (25).

General Habit Annual or perennial herbs, shrubs, or rarely trees (some species of Erycibe); often
climbing (Ipomoea) with somewhat milky sap; some with tuberous roots (Ipomoea batata) or stems,
others with rhizomes; Cuscuta is the stem parasite, twining round the host and sending haustorial
roots; some are xerophytic and thorny; others are aquatic (Ipomoea aquatica).
Leaves Simple, alternate, exstipulate, petiolate; entire or variously lobed; cordate (Ipomoea purpu-
rea), hastate (I. aquatica), deeply emarginate (I. biloba), palmately dissected (I. palmata) or pinnately
dissected (Quamoclit pennata); greatly reduced to small scales (Cuscuta).
Inflorescence Typically an axillary dichasial cyme, or flowers solitary axillary; sometimes paniculate
(Porana).
Flower Bracteate, bracts often large and showy, sometimes forming involucre; bracteolate, complete,
hermaphrodite, actinomorphic, pentamerous, hypogynous; rarely unisexual and such species are then
dioecious.
Calyx 5 sepals, usually free; persistent; quincuncial (Convolvulus, Ipomoea), or fused and valvate
(Cuscuta); 4 sepals in Hildebrandtia.
Corolla 5 united petals; often infundibuliform (Ipomoea), or salverform, or campanulate (Convolvulus);
induplicately valvate or twisted in bud; 5 coronary outgrowths are present at the base of the corolla
and alternate with the stamens in Cuscuta; 4 petals in Hildebrandtia.
Androecium 5 stamens, epipetalous, alternipetalous, inserted towards the base of corolla tube;
anthers dithecous, dorsifixed or basifixed, introrse, longitudinally dehiscent; sagittate in Ipomoea; 4
stamens in Hildebrandtia; filaments are usually of different length, i.e. heterodynamous.
Gynoecium Usually bicarpellary (rarely 3 to 5 carpels), syncarpous; superior; usually 2 locules
(rarely 4 locules formed by false septation), 1 or 2 ovules in each locule, axile placentation; style
usually 1; stigmas 1 or 2; unilocular ovary with parietal placentation is present in Erycibe: often a
cup-shaped or ring-like nectariferous disc is present below the ovary.
Fruit and Seed Fruit usually a capsule (Ipomoea), rarely a nut or berry. Seeds with large and straight
or curved embryo, and scanty and cartilaginous endosperm; sometimes hairy.
General Floral Formula Br, Brl, ≈, , K5, C (5), A5, G (2).

The family is important as a source of food, for drugs, for several ornamentals, etc.
• Plants used for edible purposes include (i) Ipomoea batata (sweet potato or Shakarkandi), of
which fleshy roots are eaten because of their rich content of sugar and starch, and are also
used for production of industrial alcohol, pectin, starch and sugar-syrup; (ii) Ipomoea aquat-
ica (Nari-ka-Sag), of which leaves and young shoots are used as vegetable, (iii) Calonyction
396 Plant Taxonomy
petal
stigma
calyx
anther stigma
corolla
lobe
filament
A Flower
style
sepal
disc connective
ovary
pedicel ovule stigma
filament
L.S. Flower
Stamen
style
flower
ovary
disc
Gynoecium
leaf
stem
Floral Diagram
Flowering Branch
Fig. 21.87 Convolvulus pluricaulis Choisy.
muricatum, of which floral pedicels are eaten, (iv) Calystegia sepium, of which roots are
cooked and eaten, (v) Rivea hypocrateriformis, of which young shoots and leaves are used
as vegetable.
• Plants of medicinal value include (i) Evolvulus alsinoides, used as a bitter tonic, (ii)
Exogonium purga, yields the drug ‘jalap’, used as a purgative, (iii) Ipomoea violacea, of
which seeds contain d-1ysergic acid amide and are hallucinogenic.
• Plants of ornamental value include (i) Argyreia speciosa (elephant creeper), Calonyction
aculeatum (moon flower), Ipomoea carica (railway creeper), I. coccinea (star Ipomoea), I.
fistulosa, I. lobata, I. purpurea (morning glory), I. quamoclit (Cyperus vine), I. tuberosa
(wood rose), I. violacea (heavenly blue), and Porana paniculata (Christmas vine).
• Plants used as sand binder are Ipomoea biloba, I. fistulosa and I. pescaprae. I. fistulosa is
also used as a hedge plant.
• Plants which become troublesome weeds are Ipomoea arvensis and Cuscuta reflexa.

The family is usually divided into 2 subfamilies, viz. Convolvuloideae (leafy autophytes; interstami-
nal scales absent) and Cuscutoideae (leafless total parasites; interstaminal scales present). However,
Bentham and Hooker divided Convolvulaceae into five tribes i.e. Convolvuleae, Dichondreae,
Nolaneae, Cresseae and Cuscuteae. Several taxonomists place Cuscuta under an independent uni-
generic family Cuscutaceae, and some also treat Dichondra and related genera in an independent
family Dichondraceae.
Convolvulaceae is related closely to Solanaceae and also to Polemoniaceae, Boraginaceae and
to some extent to Hydrophyllaceae and Nolanaceae. According to Takhtajan, it is more allied to
Polemoniaceae, Boraginaceae and Hydrophyllaceae, and thus belongs to Polemoniales. Cronquist
(1981) and Thorne (1983) treated it under Solanales.
21.75.8 Description of Common Plants

1. Convolvulus pluricaulis Choisy (Fig. 21.87)
Habit: A perennial, prostrate herb. Leaf: Simple, alternate, exstipulate. Inflorescence: Solitary axil-
lary or dichasial cyme. Flower: Bracteate, bracteolate, actinomorphic, hermaphrodite, pentamerous,
hypogynous, light purple. Calyx: 5 sepals, free, quincuncial. Corolla: 5 petals, united, infundibu-
liform, valvate. Androecium: 5 stamens, epipetalous, alternipetalous, filaments of different sizes;
dorsifixed. Gynoecium: 2 carpels, syncarpous, superior, bilocular, carpels medianly placed, 2 or
more ovules in each locule, axile placentation; stigma bifid and hairy; a nectariferous disc is present
below the ovary. Fruit: Capsule.
Floral Formula: Br, Brl, ≈, , K5, C (5), A5, G (2).
2. Cuscuta reflexa Roxb. (Amar Bel or Akash-Bel; Fig. 21.88)
Habit: A twining total parasite. Stem: Weak, twiner, pale-green, develop haustoria at the point of
contact with the host. Leaf: Absent. Inflorescence: Flowers solitary or in racemose clusters. Flower:
Bracteate, ebracteolate, hermaphrodite, actinomorphic, pentamerous, small, pale-green. Calyx: 5
sepals, fused, valvate. Corolla: 5 petals, united, campanulate, valvate, with 5 coronary outgrowths
at the base of corolla. Androecium: Same as in Convolvulus. Gynoecium: Style very much reduced,
disc red coloured; other details same as in Convolvulus. Floral Formula: Br, Ebrl, ≈, , K(5),
C (5), A5, G (2).
21.76 SOLANACEAE POTATO OR NIGHTSHADE FAMILY

398 Plant Taxonomy
stamen ovary wall

ovule
petal
locule
stigma
placenta stigmas
coronary
outgrowth T.S. Ovary
ovules
style
ovary sepal
ovary
thalamus disc
pedicel disc
L.S. Flower thalamus
pedicel
support Gynoecium
flower
stem
Flowering Branch Floral Diagram
Fig. 21.88 Cuscuta reflexa Roxb.

Herbs or shrubs, rarely trees; leaves alternate, exstipulate; flowers bisexual, actinomorphic, pentam-
erous; ovules bilocular; carpels obliquely placed; placentation axile; placenta obliquely placed; fruit
berry or capsule.

Atropa, Capsicum, Cestrum, Datura, Lycopersicon, Nicotiana, Petunia, Physalis, Solanum,
Withania.

A family of about 90 genera and 2800 species distributed in both tropical and temperate regions.
Central and South America are the chief centres of distribution where over 40 genera are found.
About 15 genera and over 90 species have been reported from India. Some of the larger genera
along with their approximately worldover reported species (Hickey and King, 1988) are Solanum
(1700), Cestrum (150), Physalis (l00), Lycium (80–90), Nicotiana (66), Capsicum (50), Petunia (40),
Hyoscyamus (20) and Datura (10). Solanum tuberosum (potato) and Nicotiana tabacum (tobacco)
are the two most utilized plants of Solanaceae.

General Habit Mostly annual (Physalis minima), biennial, or perennial herbs (Withania somnifera),
sometimes shrubs (Brunfelsia), or trees (Solanum giganteum, S. verbascifolium); rarely vines (Lycium
sinensis) or lianas.
Stem Herbaceous or woody; erect or twining, or creeping; sometimes modified into tubers (Solanum
tuberosum); often with bicollateral vascular bundles.
Leaves Usually simple, alternate, exstipulate; entire; sometimes variously dissected or lobed; oppo-
site or sub-opposite in upper part of the plant; pinnately compound in tomato and potato; unicostate
reticulate.
Inflorescence Generally cymose, or sometimes solitary flowers (Datura stramonium); sometimes
extra-axillary scorpioid cyme called rhipidium as in Solanum nigrum; or umbellate cyme (Withania
somnifera).
Flower Bracteate (Petunia) or ebracteate (Withania), pedicellate, bisexual; actinomorphic or weakly
zygomorphic due to oblique position of the ovary; pentamerous, hypogynous.
Calyx 5 sepals, gamosepalous, valvate, persistent; rarely the sepals are 4 or 6, and often enlarging
to envelope the fruit (Physalis, Withania).
Corolla 5 petals, gamopetalous, rotate to tubular (Solanum, Cestrum) or bell-shaped (Atropa)
or infundibuliform (Petunia); usually alternate with sepals; rarely bilipped and zygomorphic
(Schizanthus); usually valvate, sometimes convolute (Datura).
Androecium 5 stamens, epipetalous, alternipetalous; filaments usually of unequal length; stamens
are only 2 in Schizanthus, and 4 and didynamous in Salpiglosis; anthers dithecous; usually connivent
i.e. forming a cone by coming close together but not actually fusing (e.g. Solanum); anthers opening
either lengthwise or by terminal pores. .
Gynoecium Bicarpellary, syncarpous, carpels obliquely placed in the flower; ovary superior, bilocu-
lar, numerous anatropous or slightly amphitropous ovules in each locule; axile placentation; often
the placentae are swollen; sometimes carpels divided by a false septum; style simple, with entire or
400 Plant Taxonomy
2-lobed stigma; sometimes the carpels are more than 2 (Capsicum) and also the number of locules
is 3 to 5 (Nicandra); a hypogynous disc usually present.
Fruit and Seeds Fruit usually a berry (Physalis, Withania) or septicidal capsule (Datura). Seeds
with copious endosperm and straight or curved embryo.
Pollination and Dispersal Pollination is entomophilous. Bright and showy petals, presence of nectar
secreted by nectariferous disc and strongly scented flowers of several members of this family are the
chief attractions for insects. Flowers are protogynous and both self-pollination and cross-pollination
occur. Only self-pollination occurs in cleistogamous flowers of Salpiglossis. Dispersal of seeds takes
place mainly by birds and animals and sometimes by water (Atropa).
General Floral Formula Br or Ebr, ≈, , K(5), C (5), A5, G (2).

• Ornamental Plants: Common plants1 of ornamental value of Solanaceae are Brunfelsia
calycina (fragrant flowers), Cestrum diurnum (Day Jasmine), C. nocturnum (Night Jasmine),
Hyoscyamus niger (Henbane), Nicotiana alata (white-flowered), Petunia hybrida (pink-flow-
ered), P. violacea (white-flowered), Salpiglossis sinuata, Schizanthus pinnatus, S. retusus,
Solanum dulcamara (violet-coloured), S. jasminoides (white and blue-coloured), S. seafor-
thianum (bluish-purple coloured).
• Tobacco: Nicotiana tabacum (tobacco) leaves are dried, made into tobacco, and used
universally in cigarette, bidi, cigar, pipes, hukkah as well as for chewing and snuffing. It is
also used in medicine as a sedative and anti-spasmodic. N. rustica is also used for similar
purposes.
• Food Plants: (i) Tubers of Solanum tuberosum (potato or Aalu) are used throughout the
world as a common vegetable and also used for the production of starch, dextrin, several
alcohols and some other industrial products, (ii) fruits of Lycopersicon lycopersicum syn.
Solanum lycopersicum are the tomato (Tamatar), used as a delicious vegetable and eaten
raw, (iii) fruits of Solanum melongena (Brinjal, eggplant or Baingan) are eaten as vegetable,
(iv) chillies, ‘Mirch’ or red pepper are the fruits of Capsicum annuum and C. frutescens,
(v) Physalis peruviana (Raspberry or Cape Gooseberry) yield the delicious edible fruits.
• Medicinal Plants2 : (i) Atropa belladona (Belladona or deadly nightshade) roots yield a
powerful alkaloid ‘atropine’ used in belladona plasters, tinctures etc., for relieving pain,
and also for dilating pupils of eyes for eye-testing, (ii) Datura stramonium (Datura) leaves
and flowers are the source of the drug “stramonium”, used in asthama, and its seeds are
deadly poisonous, (iii) Hyoscyamus niger (Henbane) leaves yield the alkaloid hyoscyamine
used in treating asthama and whooping cough, (iv) Mandragora autumnalis roots are used
as sedative and hypnotic, (v) Nicotiana tabacum leaves are used as sedative, antispasmodic
and also in iradicating animal pests, such as lices, (vi) Solanum surattense roots are used in
1
2
asthma and leaves in rheumatism, (vii) Withania coagulans fruits are used in curing asthma
and liver troubles, (viii) Withania somnifera (Asgandh) roots are used in curing cough and
rheumatism.

Variously placed under Polemoniales (Bentham and Hooker), Solanales (Hutchinson, Cronquist,
Thorne) and Scrophulariales (Takhtajan), the family Solanaceae has been divided by Bentham and
Hooker into 5 tribes (Solaneae, Atropeae, Hyoscyameae, Cestrineae and Salpiglossideae). Wettstein
divided Solanaceae into 5 different tribes namely Nicandreae, Solaneae, Datureae, Cestreae and
Salpiglossideae.
Solanaceae is allied closely to Convolvulaceae in both having pentamerous flowers, persistent calyx
and bicarpellary ovary. However, Solanaceae possess obliquely placed ovary, and are predominantly
erect whereas these characters are absent in Convolvulaceae. In certain characters Solanaceae is
closely allied to Scrophulariaceae, Boraginaceae and Polemoniaceae. Hutchinson has raised the tribe
Salpiglossideae to the rank of an independent family Salpiglossidaceae.

Solanum nigrum L. (Black Nightshade or Makoi, Fig. 21.89)
Habit: A small annual herb. Leaf: Simple, alternate but opposite in the floral region; petiolate,
exstipulate, ovate, entire or slightly lobed, acute, unicostate reticulate. Inflorescence: Extra-axillary
scorpioid cyme called rhipidium. Flower: Ebracteate, pedicellate, hermaphrodite, actinomorphic, pen-
tamerous, hypogynous, white. Calyx: 5 sepals, fused, valvate, persistent. Corolla: 5 petals, gamopeta-
lous, valvate. Androecium: 5 stamens, free, epipetalous, filaments short; anthers conniving and form-
ing an envelope around the style, dithecous, basifixed, with apical pores. Gynoecium: Bicarpellary,
syncarpous, superior, bilocular, many ovules in each locule, axile placentation, septum oblique and
placentae are highly swollen; style long and hairy at the base; stigma bifid. Fruit: Berry.
Floral Formula: Ebr, ≈, , K(5), C (5), A5, G (2).
21.77 POLEMONIACEAE PHLOX FAMILY


Milky latex absent; flowers pentamerous; calyx gamosepalous; stamens inserted at different levels
on corolla tube; ovary typically tricarpellate; ovules and seeds numerous.

Phlox, Polemonium, Gilia.
402 Plant Taxonomy
style stigma
stigma
style
anther lobe
filament anther
lobe
petal
pore petal
sepal
ovary anther lobe
thalamus pedicel pedicel

L.S. Flower connective A Flower
filament
flower
Stamen
leaf
fruit
stem
Floral Diagram
Flowering Branch
Fig. 21.89 Solanum nigrum L.

A family of about 18 genera and over 300 species, Polemoniaceae are distributed chiefly in North
America, some in South America, Europe and North Asia. Some of the large genera along with the
approximate number of their reported species are Gilia (120), Phlox (77), Polemonium (52), Cobaea
(18) and Collomia (15). Colourful corollas of Polemoniaceae, specially of Phlox, have attracted the
attention of many and are thus adopted into the gardens throughout the world.

General Habit Annual or perennial herbs, rarely shrubs, small trees, or twining vines; milky latex
or coloured sap absent.
Stem Erect, hollow, angled (Polemonium caeruleum, Fig. 21.90).
flowers
F
A Fruit
ovules
loculi
leaflets D
T.S. Ovary
petal
leaf stem
sepal
A ovary (hairy)
A Flowering Branch (superior)
ovules
C
L.S. Ovary
stigma
petals
sepal stamen
(hairy)
stigma
style
style
E
Style and Stigma
ovary B
A Bisexual Flower
Fig. 21.90 Polemonium caeruleum L.
404 Plant Taxonomy
Leaves Alternate or opposite; simple or compound; exstipulate; stipules, if present are large and
foliaceous (Cobaea); in Polemonium caeruleum the leaves are pinnate, lower petiolate while the
upper ones are sessile; 6–12 pairs of lateral leaflets are lanceolate to oblong in shape with acuminate
apex (Fig. 21.90).
Inflorescence Usually cymose, corymbose to capitate; rarely flowers solitary and axillary.
Flowers Bracteate, bracteolate, complete, hermaphrodite; actinomorphic, rarely slightly zygomorphic
(Bonplandia); pedicellate, pentamerous and usually showy.
Calyx Sepals 5, gamosepalous; valvate or imbricate; persistent.
Corolla Petals 5, gamopetalous, campanulate (Cobaea), funnel-shaped or salver-shaped; usually
convolute; corolla tube usually well-developed.
Androecium Stamens 5, epipetalous and alternipetalous; attached on the corolla tube at various
heights; anthers dithecous; an intrastaminal disc usually present.
Gynoecium A compound pistil of usually 3 united carpels; trilocular, one to numerous ovules in
each locule; axile placentation; ovary superior; style, filiform; stigmas 3 or rarely 2.
Fruit Usually a loculicidal capsule; sometimes septicidal (Cobaea); rarely indehiscent.
Seeds Usually with abundant and fleshy endosperm; in Cobaea, however, seeds have no endosperm;
seeds sometimes covered by mucilaginous coat (Collomia).

Polemoniaceae is not of much economic importance except that of a few ornamentals grown as
annuals, biennials, perenniales or vines. Some of such plants of ornamental value belong to Phlox,
Polemonium, Cobaea, Gilia and Linanthus. Beautiful flowers come from Phlox drummondii
(annual), P. paniculata (perennial) and Cobaea scandens (climber), etc.

Bentham and Hooker placed Polemoniaceae and four more families (Boraginaceae, Hydrophyllaceae,
Convolvulaceae and Solanaceae) under order Polemoniales of series Bicarpellatae of subclass
Gamopetalae. Along with 18 more families, all these five families have been placed under order
Tubiflorae by Engler and Prantl. Hutchinson included only 3 families (Polemoniaceae, Cuscutaceae
and Hydrophyllaceae) under order Polemoniales of division Herbaceae.
Phylogeny of Polemoniaceae has been perplexing botanists since long. According to Hallier,
Polemoniaceae has been “probably derived from Linaceae”. Bessey considered it to be derived from
Boraginaceous stock whereas Rendle believed Polemoniaceae to have been derived from sympeta-
lous relatives of Rosales. According to Hutchinson, however, Polemoniales have been derived from
Geraniales. Some morphologists, however, believe that Polemoniaceae are closely related to both
Caryophyllaceae and Geraniaceae (Dawson, 1936).
21.78 PERSONALES
1. Mostly herbs or undershrubs, rarely trees.
2. Leaves are exstipulate.
3. Flowers zygomorphic and hypogynous.
4. Number of stamens are generally less than corolla lobes, and epipetalous.
5. Stamens are usually didynamous.
6. Internal phloem is present in members of some families.
7. Fruit usually a capsule.
Bentham and Hooker included 8 families under Personales. These are Scrophulariaceae,
Orobanchaceae, Lentibulariaceae, Bignoniaceae, Pedaliaceae, Columelliaceae, Gesneriaceae and
Acanthaceae. Engler and Diels discussed all these families under order Tubiflorae.
Only Scrophulariaceae, Bignoniaceae, Pedaliaceae and Acanthaceae have been discussed in this
text.
21.79 SCROPHULARIACEAE SNAPDRAGON FAMILY

Gamopetalae, Bicarpellatae, Personales.

Mostly herbs or shrubs, rarely trees; flowers pentamerous; zygomorphic, corolla bilipped; usually
with 2 or 4 fertile stamens, epipetalous; ovary bilocular, superior.

Antirrhinum, Digitalis, Linaria, Lindenbergia, Mazus, Verbascum, Veronica.

Represented by about 220 genera and 3500 species (Hickey and King, 1988), Scrophulariaceae are
cosmopolitan in their distribution. About 60 genera and 360 species have been reported from India,
chiefly from the Himalayas. Some of the genera along with their approximately reported species
from worldover are Pedicularis (500), Calceolaria (300–400), Scrophularia (300), Verbascum
(300), Veronica (300), Penstemon (280), Linaria (150), Castilleja (150), Selago (150), Sutera (130),
Mimulus (100), Antirrhinum (40) and Digitalis (20). The common names of the family (snapdragon,
foxglove or figwort family) are because of its genera Antirrhinum (snapdragons), Digitalis (foxglove)
and Scrophularia (figwort).
406 Plant Taxonomy

General Habit Mostly herbs or undershrubs, and only a few are big shrubs (some species of
Veronica) or trees (Paulownia); several are climbers (Maurandia), and some are saprophytic or
parasitic (Hyobanche); Lathrea is a chlorophyll-less root parasite while Euphrasia and Pedicularis
are semi-parasitic.
Leaf Simple or rarely pinnately compound; exstipulate; alternate (Verbascum), opposite (Mimulus)
or whorled (Russelia); Ambulia shows heterophylly; both stem and leaves are generally hairy.
Inflorescence Variable from racemose (Mazus) or cymose (Verbascum), or flowers solitary axillary
(Linaria, Lindenbergia); spike (Digitalis), panicle (Scrophularia himalayensis).
Flower Bracteate, bracteolate, bisexual; zygomorphic but sometimes nearly actinomorphic
(Verbascum); upper leaves and bracts are sometimes brightly coloured (Castilleja); hypogynous;
usually with an annular disc.
Calyx Usually 5 united sepals; imbricate, quincuncial or valvate; sepals are 4 in Veronica and
Scoparia; sometimes 2 anterior sepals are united (Calceolaria).
Corolla Usually 5 united petals; often bilipped (2/3); sometimes only 4 petals are present, and the
one placed anteriorly is smaller than the others (Veronica, Fig. 21.91A); frequently the anterior petals
are spurred (Linaria) or with a gibbous (Antirrhinum, Fig. 21.91B) to saccate base; in Verbascum
corolla is regular with all the 5 equal petals (Fig. 21.91C).
Androecium 4 stamens (Mazus), sometimes only 2 stamens (Veronica); rarely all the 5 stamens are
fertile (Verbascum); didynamous, epipetalous; sometimes the posterior stamen is represented by a
staminode (Antirrhinum, Scrophularia, Penstemon); fertile stamens are alternate with corolla lobes;
anthers dithecous, occasionally connivent, dehiscence mostly longitudinal and rarely poricidal.
Gynoecium Bicarpellary, syncarpous, superior, bilocular, numerous anatropous or amphitropous
ovules in each locule, axile placentation; carpels usually anterio-posteriorly placed; style simple;
stigma bilobed; nectar-secreting disc present below this ovary.
Fruit and Seeds Fruit usually a capsule, rarely a berry (Leucocarpus) or drupe surrounded by per-
sistent calyx. Seeds numerous, small, endospermic with straight or slightly curved embryo.
Pollination and Dispersal Flowers usually adapted to insect-pollination. Insects are attracted because
of coloured corolla and nectar-secreting hypogynous disc. Flowers which have long-tubed corolla
(e.g. Scrophularia) are pollinated by wasps while those having short-tubed corolla (e.g. Veronica,
Verbascum) are pollinated by bees. Dispersal of seeds usually takes place by water, birds or
animals.
General Floral Formula Br, Brl, , , K(5), C (5), A4 or 2, G (2).

• Ornamental plants1 of the family include Angelonia angustifolia, Antirrhinum majus (snap-
dragon), Calceolaria scabiosaefolia (slipperwort), Castilleja (Indian paintbrush), Chelone
1
A C
B
Fig. 21.91A–C Floral diagrams of Veronica anagallis (A), Antirrhinum orontium (B) and Verbascum
chinense (C).
glabra (turtlehead), Collinsia grandiflora, Digitalis purpurea (foxglove), Linaria vulgaris

(toadflax), Mimulus luteus (monkey flower), Nemesia versicolor (Nemesia), Paulownia
tomentosa, Pedicularis canadensis (housewort), Russelia juncea (coral blow), Torenia asi-
atica and Veronica bungifolia (speedwell).
• Medicinal plants1 of the family include (i) Bacopa monnieri (used in epilepsy and insan-
ity), (ii) Digitalis purpurea (dried leaves are used in congested heart failure), (iii) Herpestis
cuneifolia (used in rheumatism), (iv) Picrorhiza kurroa (used as a laxative and cathartic),
(v) Scoparia dulcis (used in fever, cough and bronchitis), and (vi) Verbascum sinensis (used
in dysentery).

Scrophulariaceae has been variously placed under Personales (Bentham and Hooker, and Hutchinson),
Scrophulariales (Takhtajan, Cronquist) and Bignoniales (Thorne). Wettstein divided Scrophulariaceae
into 3 subfamilies viz. Verbascoideae, Scrophularioideae and Rhinanthoideae.
1
408 Plant Taxonomy
Scrophulariaceae is closely allied to Solanaceae and Convolvulaceae having exstipulate leaves, per-
sistent gamosepalous calyx and bicarpellary superior ovary. It is also closely related to Bignoniaceae
having zygomorphic flowers, exstipulate leaves and hypogynous disc. In several characters
Scrophulariaceae is closely allied also to Labiatae and Acanthaceae.

1. Mazus japonicus (Thunb.) Kuntze syn. M. rugosus Lour. (Fig. 21.92)
Habit: Small annual herb. Leaf: Radical, simple, exstipulate, sessile but lamina narrowed basally into
the petiole, spathulate, dentate. Inflorescence: Racemose raceme raised on a scape. Flower: Bracteate,
pedicellate, complete, hermaphrodite, zygomorphic, bilipped, pentamerous, hypogynous. Calyx: 5
sepals, poly- or gamosepalous, valvate or quincuncial. Corolla: 5 petals, united; bilabiate (2/3) person-
ate, posterior lip bilobed while anterior lip trilobed. Androecium: 4 stamens, polyandrous, epipetalous,
stigma stigma
anther corolla
filament style style
ovary ovule
calyx
sepal
hairs A Flower
pedicel flower
L.S. Flower ovary
Gynoecium
scape
leaf
roots
Floral Diagram Flowering Plant
Fig. 21.92 Mazus japonicus Thunb.

didynamous, filaments curved apically; anthers dithecous, basifixed, introrse. Gynoecium: Same as
in family description. Fruit: Capsule.
Floral Formula: Br, Ebrl, , , K5, C (2/3) A2+2, G (2).
21.80 BIGNONIACEAE BIGNONIA FAMILY


Woody lianas, shrubs or trees; leaves usually opposite and compound; inflorescence cymose; flowers
zygomorphic; fruit capsule.

Jacaranda, Kigelia, Millingtonia, Tecoma, Heterophragma, Oroxylum and Dolichandrone.

Represented by about 120 genera and over 650 species, Bignoniaceae are distributed mainly in tropi-
cal and subtropical, and a few in temperate regions. About 15 genera and 40 species are found in
India. Some of the larger genera along with their approximate number of worldover reported species
are Tabebuia (100), Arrabidaea (70), Jacaranda (50), Anemopaegma (30), Tecoma (16), Catalpa
(11) and Crescentia (5).

General Habit Mostly woody lianas or climbers, climbing with the help of tendrils; or trees or
shrubs; only rarely herbs; some climb with the help of aerial roots (Campis radicans), others are
twiners (Pandorea, Tecomaria) and most genera are tendril climbers.
Anatomically, wedges of phloem develop in stem in between secondary xylem.
Leaf Usually opposite, pinnately compound; exstipulate; terminal leaflet usually modifying into
tendril; tendrils may be simple or branched, hooked, or provided with some adhesive discs; some-
times the leaves are simple and rarely whorled (Diplanthera).
Inflorescence Usually cymose (dichasial cyme) with bracts and bracteoles.
Flower Bracteate, bracteolate, complete, hermaphrodite, zygomorphic, hypogynous, often showy.
Calyx Usually 5 united sepals, 5-toothed, often campanulate, valvate.
Corolla Usually 5 united petals, campanulate or funnel-shaped, imbricate; sometimes bilipped (2/3)
with the upper lip of 2 petals and lower lip of 3 petals.
Androecium Typically 4, epipetalous stamens, didynamous, alternipetalous, with a posterior stamin-
ode; sometimes 2 (Catalpa, Fig. 21.93) and rarely 5 (Oroxylum) stamens; usually the anther cells
one above the other; dithecous, introrse, longitudinal dehiscence.
410 Plant Taxonomy
stigma
style
sepal
staminode
stamen stamen
ovary
L.S. Flower
corolla tube Stamen
corolla tube
Part of the
sepal Corolla Tube style
larger lip
ovules
ovary
flower
A Flower disc
calyx
stigma L.S. Ovary
style
T.S. Ovary
Style
(upper portion)
flower
stem
leaf
Flowering Branch
Fig. 21.93 Catalpa bignonioides Walt.

Gynoecium Bicarpellary, syncarpous, superior, bilocular, numerous anatropous ovules in each locule,
axile placentation; sometimes unilocular (Kigelia) with parietal placentation; style simple, terminal;
stigma bilipped; a hypogynous nectar-secreting disc is present.
Fruit and Seeds Fruit usually a capsule, sometimes fleshy and indehiscent. Seeds often winged,
non-endospermic, with straight embryo.
General Floral Formula Br, Brl, , , K(5), C (5), A4+1 staminode , G (2).

• Ornamental plants of the family include handsome trees (Catalpa bignonioides, Haplophragma
adenophyllum, Jacaranda mimosaefolia, Kigelia pinnata, Millingtonia hortensis, Spathodea
campanulata and Tabebuia spectabilis), shrubs (Bignonia unguis-cati), beautiful tendril-
climbers (Adenocalymna alliaceum, Bignonia venusta syn. Pyrostegia venusta), aerial rootlet
climbers (Campis grandiflora and C. radicans) and herbaceous genus Incarvillea.
• Some of the timber-yielding plants of the family include species of Catalpa (Fig. 21.93),
Dolichandrone, Haplophragma, Millingtonia, Oroxylum, Radermachera, Spathodea,
Stereospermum, Tabebuia and Tecoma.
• An inferior quality of cork is prepared from the bark of Millingtonia hortensis (Akash Neem
or Indian cork tree).
• Crescentia cujete (Calabash-tree) is grown for its gourd-like fruits, which, when hollowed
out and dried, are used for holding liquids.

Variously placed under Personales (Bentham and Hooker), Bignoniales (Hutchinson, Thorne) and
Scrophulariales (Takhtajan, Cronquist), the family Bignoniaceae has been divided into 4 tribes
(Bignonieae, Tecomeae, Eccremocarpeae and Crescentieae) by Schumann in Engler and Prantl’s
Pflanzenfamilien. In possessing didynamous stamens, bicarpellary ovary, zygomorphic flowers and
capsular fruits, Bignoniaceae is closely allied to Scrophulariaceae. In some respects it is also allied
to Acanthaceae and Pedaliaceae.
21.81 PEDALIACEAE BENNE FAMILY


Herbs, rarely shrubs; leaves opposite or the upper ones alternate; flowers usually solitary axillary;
flowers bisexual, zygomorphic; calyx and corolla 5 and united; stamens generally 4 and didynamous;
a small staminode present; fruit capsule, often prickly.
412 Plant Taxonomy

Sesamum (Fig. 21.94), Martynia, Pedalium.

Represented by about 15 genera and about 65 species, the members of this family are distributed
largely on sea shores and desert areas occurring in Indo-Malaysian regions, Madagascar, South Africa
and tropical Australia. In India, the family is represented by three genera, occurring chiefly in south
Indian states. Sesamum indicum (Sesame or Gingelly) is cultivated as an oil-seed crop throughout
the warmer parts of India.

General Habit Annual or perennial herbs or undershrubs, with mucilage containing glandular hairs;
sometimes roots are tuberous (Martynia).
Leaf Opposite or spirally arranged, sometimes the upper leaves alternate; simple, exstipulate; entire,
lobed or divided.
Inflorescence Usually solitary axillary flowers, sometimes in three-flowered axillary cymes; some-
times terminally racemose (Martynia).
Flower Bisexual, zygomorphic, hypogynous; characteristic glands are present at the base of floral
stalk; these are actually metamorphosed flowers.
Calyx Usually 5 sepals, more or less united; rarely sepals distinct (Martynia).
Corolla Petals 5, united; tubular-ventricose; limb obscurely bilabiate, 5-lobed and the lobes
imbricate.
Androecium Stamens 4, didynamous, epipetalous on corolla tube; small posterior staminode rep-
resents fifth stamen; two fertile stamens and remaining three staminodes are present in Martynia;
anthers dithecous, introrse and dehisce longitudinally.
Gynoecium Bicarpellary, syncarpous, superior; ovary bilocular or becomes tetralocular due to the
formation of false septa; one to many ovules in each locule; axile placentation; ovary sometimes
unilocular with two parietal placentae in Martynia; style one, slender; stigma bilobed; at the base
of ovary is present a nectar-secreting disc.
Fruit and Seed Fruit capsule or sometimes nut, commonly spiny; two long horns are present in
Martynia. Seeds are non-endospermic or with a thin endosperm possessing a straight embryo.
General Floral Formula EBr, , , K(5), C (5), A4+1 staminode, G (2).

• “Sesame oil”, a well-known edible oil, is extracted from the seeds of Sesamum indicum
(Sesame or Til; Fig. 21.94). It is widely used in confectionary, in the manufacture of several
cosmetics, insecticides and soaps and also for making margarine. Hindus use the seeds in
flowers
anther
lobes
filament D
Stamen
fruit
G
A Fruit
leaf
seeds
stem
A
A Flowering Branch
B F H
L.S. Ovary L.S. Fruit
A Flower
corolla
gynoecium
calyx
stamens C
I
Gynoecium Corolla and Stamens
Floral Diagram (with expanded calyx) (expanded)
Fig. 21.94 Sesamum indicum L.
many of their religious ceremonies. Seeds are widely used as nourishing food and also as
flavouring agents. Oil is also extracted from Sesamum angustifolium and used for similar
purposes.
• Leaves of Pedalium murex and Ceratotheca sesamoides are used as vegetables.

Family Pedaliaceae has been placed differently under order Personales by Bentham and Hooker,
Tubiflorae by Engler and Prantl, Scrophulariales by Takhtajan (1980) and Cronquist (1981) and under
414 Plant Taxonomy
order Bignoniales by Hutchinson (1973) and Thorne (1983). Bentham and Hooker (1862) divided
family Pedaliaceae into four tribes namely Martynieae, Pedalieae, Sesameae and Pretreae. Many
botanists treat the tribe Martynieae as a separate family, Martyniaceae, due to the presence of char-
acters such as unilocular ovary, parietal placentation and horned capsules. Members of Pedaliaceae
resemble quite closely to Bignoniaceae. Some taxonomists also treat Martynia and other related
genera in family Bignoniaceae.
As far as the chemical features are concerned, saponins are lacking in all members of family
except Sesamum indicum. Tannins are absent in S. indicum and Pedalium murex while they are doubt-
fully present in Sesamum laciniatum and Martynia annua. Steroids are present in S. indicum and S.
laciniatum while they are absent in Pedalium murex and Martynia annua. Leucoanthocyanins are
present in Sesamum laciniatum and Martynia annua while they are absent in Sesamum indicum and
Pedalium murex. All these facts indicate that Martynia annua resembles other taxa of Pedaliaceae,
hence its inclusion in the family seems correct. Das et al. (1985), however, opined to treat Martynia
in an independent family Martyniaceae on the basis of their studies of the distributional pattern of
phenolic acids.
21.82 ACANTHACEAE ACANTHUS FAMILY


Herbs, shrubs or trees; leaves opposite decussate; flowers zygomorphic; corolla bilipped (2/3); seeds
with jaculators.

Acanthus, Adhatoda, Barleria, Eranthemum, Justicia, Peristrophe, Ruellia, Thunbergia.

A family of over 250 genera and 2500 species, Acanthaceae are distributed mainly in tropics, but
also in Mediterranean region, Australia and USA. Central America, Brazil, Africa and Indo-Malaysia
are the 4 chief centres of distribution. About 70 genera and over 340 species have been reported
from India. Some of the largely represented genera along with the approximate number of their
worldover reported species are Justicia (300), Strobilanthes (250), Barleria (230), Thunbergia (200),
Aphelandra (200), Decliptera (150), Blepharis (100), Staurogyne (80), Mendoncia (60), Crossandra
(60), and Acanthus (50).

General Habit Mostly perennial herbs or shrubs, rarely trees; some are twiners (Thunbergia,
Mendoncia), or xerophytes (Barleria, Blepharis), or halophytes (Acanthus ilicifolius), or spiny shrubs
(Acanthus montanus).
Leaf Simple, opposite decussate, exstipulate, entire, rarely pinnately lobed with small spines on the
margins (Acanthus mollis); cystoliths, appearing as protuberances or streaks, are usually common
on stems and leaves.
Inflorescence Usually a dichasial cyme, frequently condensed in the leaf axils; sometimes racemose
spike (Adhatoda vasica), or flowers solitary axillary (Thunbergia).
Flower Usually bracteate, bracteolate, pedicellate or subsessile, complete, hermaphrodite, zygomor-
phic, hypogynous; bracteoles are often large, coloured and more or less enclosing the flower; in
Peristrophe (Fig. 21.95) out of the 2 bracts the posterior one is larger than the anterior one, and 4
laterally placed bracteoles cover the calyx.
Calyx Usually 5 united (Justicia) or free (Adhatoda) sepals; valvate (Justicia, Fig. 21.96) or quin-
cuncial (Adhatoda); sometimes sepals are 4 or rarely 3; rarely the calyx is much reduced and divided
into teeth-like structures (Thunbergia).
Corolla Usually 5 lobed, gamopetalous, imbricate or twisted; bilipped (2/3) or bilabiate personate,
of which the upper lip is bifid or bilobed and the lower lip is trilobed; sometimes the upper lip is
not developed (Acanthus); corolla is tubular in Thunbergia and Ruellia.
Androecium Usually 4 didynamous stamens (Ruellia, Thunbergia), or only 2 stamens (Adhatoda,
Justicia, Peristrophe), epipetalous, usually exserted; rarely 5 stamens (Pentstemonacanthus); fre-
quently 1–3 staminodes present; anthers bilobed with often one smaller lobe than the other, connec-
tive often long (Fig. 21.95); introrse, longitudinal dehiscence.
Gynoecium Bicarpellary, syncarpous; carpels median; ovary superior, bilocular, one to many ana-
tropous ovules in each locule, axile placentation; style narrow and long; stigmas 2, of which the
anterior one is often larger; usually with a hypogynous, nectar-secreting disc.
Fruit and Seeds Fruit usually a capsule, loculicidal to the very base; rarely a drupe (Mendoncia).
Seeds 1 to many, of which the funiculus develops into a hook-like retinaculum or “jaculator”; often
non-endospermic.
Pollination Flowers are suited for insect pollination because of coloured bilabiate corolla and abun-
dant nectar in hypogynous disc. Protandry favours cross-pollination.
General Floral Formula Br, Brl, , , K(5) or 5, C (2/3), A2 or 4, G (2).

• Ornamental plants include Aphelandra aurantiaca, Barleria cristata, B. gibsoni, B. lupulina,
B. montana, Beloperone guttata (shrimp plant), Crossandra infundibuliformis, Eranthemum
bicolor, E. nervosum, Fittonia gigantea, Jacobinia carnea, J. tinctoria, Justicia gendarussa,
Ruellia brittoniana, R. tuberosa, Strobilanthes dyerianus, S. isophyllus, Thunbergia coccinia,
T. erecta, T. fragrans and T. grandiflora.
• Plants of medicinal value include (i) Adhatoda vasica (Vasaka or Basak)–leaves are use-
ful in cough, asthma and bronchitis, (ii) Asteracantha longifolia–seeds and roots are used
against cough, (iii) Barleria prionitis–leaves and roots are used against bronchitis and
cough, (iv) Barleria cristata–seeds are used as an antidote for snakebite, (v) Hygrophila
spinosa-leaves and roots are used against jaundice and rheumatism, (vi) Peristrophe
416 Plant Taxonomy
anther
lobes stigma
corolla
stigma
style
stamen
anther
lobe
style connective
bracteole A Flower
ovary
disc
ovules sepal
filament pedicel
ovary disc Gynoecium
thalamus pedicel Stamen
L.S. Flower
flower
Floral Diagram
leaf
stem
Flowering Branch
Fig. 21.95 Peristrophe bicalyculata Nees.
bicalyculata—leaves and roots are used against snakebite, (vii) Ruellia prostrata—leaf
juice is used in earache.
• A blue dye is obtained from the leaves of Strobilanthes cusia.

Acanthaceae is varyingly placed under Personales (Bentham and Hooker, and Hutchinson), Tubiflorae
(Engler and Prantl) and Scrophulariales (Takhtajan, Cronquist). It is usually divided into 4 subfamilies
(Nelsonioideae, Mendoncioideae, Thunbergioideae and Acanthoideae).
Subfamily Nelsonioideae is very close to family Scrophulariaceae, and its genera are included
in that family by several workers. Subfamilies Mendoncioideae and Thunbergioideae are interme-
diate between Bignoniaceae and Acanthaceae and are usually considered as independent families
Acanthaceae is considered to have been derived from Scrophulariaceae.

Justicia gendarussa Burm. (Fig. 21.96)
Habit: A perennial herb. Leaf: Simple, stipulate, opposite decussate, subsessile, lanceolate, obtuse,
unicostate reticulate. Inflorescence: Dichasial cyme arranged in racemose manner. Flower: Bracteate,
bracteolate, pedicellate, complete, zygomorphic, pentamerous, hypogynous. Calyx: 5 sepals, united,
valvate. Corolla: 5 petals, united; 2/3 bilabiate personate, of which posterior lip is bilobed while
anterior lip is trilobed; valvate. Androecium: 2 stamens, polyandrous, epipetalous, anther lobes situ-
ated at unequal heights; dithecous, basifixed, introrse. Gynoecium: Same as in description of the
family. Fruit: Capsule.
Floral Formula: Br, Brl, , , K(5), C (2/3), A2, G (2).
21.83 LAMIALES
1. Mostly herbs, some are shrubs and only rarely trees.
2. Leaves are generally simple, opposite or rarely whorled, and exstipulate.
3. Flowers mostly zygomorphic and pentamerous; bilabiate in Labiatae.
4. Gynoecium bicarpellary, syncarpous, superior; generally deeply 4-loobed.
5. Placentation usually axile.
6. Style gynobasic.
7. Fruit schizocarpic.
8. Seeds with scanty or no endosperm and straight embryo.
Bentham and Hooker included 4 families (Labiatae, Verbenaceae, Selaginaceae and Myoporaceae)
under Lamiales. Engler and Diels, however, included all these families, except Selaginaceae under
order Tubiflorae. Actually, it was Bessey who separated Labiatae and Verbenaceae as of a distinct
order, the Lamiales, on the basis of corolla zygomorphy and gynoecial characters.
Only Labiatae and Verbenaceae of Lamiales have been discussed in this text.
418 Plant Taxonomy
petal
stamen
stigma
style
sepal
ovary
Floral Diagram
pedicel
L.S. Flower
ovary wall
ovule
inflorescence flower locule
placenta
T.S. Ovary
anther
lobe
connective
leaf
filament
stem
Flowering Branch A Stamen
Fig. 21.96 Justicia gendarussa Burm.
21.84 LABIATAE OR LAMIACEAE MINT FAMILY

Gamopetalae, Bicarpellatae, Lamiales.

Aromatic herbs or shrubs with square stems; leaves opposite decussate; inflorescence verticillaster;
flowers zygomorphic and pentamerous; ovary deeply 4-lobed; style gynobasic; fruit schizocarpic.

Coleus, Lamium, Lavandula, Leucas, Mentha, Ocimum, Salvia.

A family of about 200 genera and 3500 species, Labiatae are cosmopolitan in their distribution
and their chief centre is the Mediterranean region. About 65 genera and over 400 species have
been reported from India, mainly distributed in North Western India and South India. Some largely
represented genera along with the approximate number of their worldover reported species and
common names of some of them are Salvia (500–700, sage), Hyptis (400), Thymus (300–400,
thyme), Teucrium (300, germander), Stachys (300), Scutellaria (200, skullcap), Coleus (200, Jacob’s
coat), Ocimum (150, basil), Nepeta (150, catnip), Lavandula (28, lavender), Mentha (25, mint) and
Rosmarinus (3, rosemary).

General Habit Usually perennial or annual terrestrial herbs with square stems, sometimes shrubs
(Orthosiphon), rarely trees (Hyptis, Leucosceptrium), or vines (Scutellaria); plant parts usually cov-
ered with glandular hairs; usually with aromatic oils.
Stem Usually quadrangular and covered with glandular hairs.
Leaf Usually simple, opposite, exstipulate; often hairy and with epidermal glands secreting volatile
oils; sometimes whorled (Dysophyla); rarely pinnately dissected or compound.
Inflorescence Often cymose, the cymes at the nodes condensed into a false whorl or verticillaster;
rarely flowers are in simple racemes (Scutellaria), or solitary and axillary; very rarely in heads
(Hyptis).
Flower Usually bracteate, bracteolate, complete, hermaphrodite, zygomorphic, hypogynous; rarely
nearly actinomorphic (some species of Mentha).
Calyx 5 sepals, gamosepalous or united into a funnel-shaped or campanulate tube; tube often ribbed;
persistent in fruit; valvate or imbricate; usually 2-lipped (1/4 in Ocimum and 3/2 in Salvia).
Corolla 5 united petals; 2-lipped (4/1 in Ocimum i.e. 4 in posterior lip and 1 in anterior lip, or
2/3 in Salvia, Leucas); corolla is usually differentiated into a tube and a limb; valvate (Ocimum),
imbricate (Salvia) or contorted.
420 Plant Taxonomy
Androecium Usually 4 stamens (Ocimum), didynamous, epipetalous; sometimes only 2 stamens

(Salvia), in each of which the fertile anther lobe is attached at the anterior end of very long con-
nective while at the posterior end is present the sterile anther lobe (Fig. 21.97); filament is usually
free and rarely connate (Coleus); dithecous, dorsifixed, introrse; dehiscence longitudinal; usually the
missing stamens are present in the form of staminodes.
Gynoecium Bicarpellary, syncarpous; ovary superior, deeply 4-lobed; bilocular when young but
becomes quadrilocular due to the formation of false septum; 2 ovules in each locule in bilocular
condition, but in quadrilocular condition one anatropous ovule is present in each locule; axile pla-
centation; style gynobasic; stigma bifid; a 4-lobed, hypogynous, nectar-secreting disc is present.
Fruit and Seeds Fruit schizocarp of 1 to 4 nutlets, each nutlet containing one seed; rarely a drupe.
Seeds with scanty or no endosperm and straight embryo.
connective style petal

stigma
filament stigma
A Stamen stamen
ovule
style pedicel ovary

calyx
nectary
L.S. Flower
ovary
nectary
Gynoecium
flower
leaf
stem
Fig. 21.97 Salvia officinalis L.

Pollination Pollination is entomophilous. Coloured bracts (Salvia), corolla and nectar in several other
genera attract the insects. Usually the lower lip of the corolla forms the platform for the visiting
insects. The long connective in Salvia also proves helpful in shedding pollens from the fertile anther
lobe. Cross-pollination is effected because the flowers are usually protandrous. Long-tubed flowers
of Monarda and several other genera are pollinated by butterflies and moths.
General Floral Formula Br, Brl, , , K(5) or (1/4) or (3/2), C (4/1) or (2/3), A2 or 4 or 2+2 , G (2).

• Garden Ornamentals: Coleus blumei, C. thyrsoideus, Lavandula officinalis, Ocimum
basilicum (sweet basil), Rosmarinus officinalis (rosemary), Salvia coccinea, S. officinalis
(garden sage), S. splendens (scarlet sage), and Thymus vulgaris (thyme).
• Volatile Oils:
(i) Basil oil, obtained from Ocimum basilicum (sweet basil), O. canum (Ram Tulsi) and
O. sanctum (holy basil or Tulsi), is used in variety of medicines and perfumery.
(ii) Lavender oil (Lavandula latifolia, L. spica, L. vera) is used in perfumery and
cosmetics.
(iii) Mint oil (Mentha arvensis, M. longifolia, M. spicata) is used in nausia and vomiting.
(iv) Origanum oil (Origanum vulgare) is used in cosmetics and soap industry.
(v) Patchouli oil (Pogostemon benghalense, P. cablin) is used in perfumery.
(vi) Pennyroyal oil (Mentha pulegium) is used in preparing synthetic menthol.
(vii) Peppermint oil (Mentha piperata) is used in pharmacy and confectionery.
(viii) Perilla oil (Perilla frutescens) is used in paints, varnishes and printing ink.
(ix) Rosemary oil (Rosmarinus officinalis) is used in medicines and perfumery.
(x) Sage oil (Salvia splendens) is used in soap industry.
(xi) Sweet Majoran oil (Majorana hortensis) is used in perfumery and liquor industry.
• Medicinal Plants: (i) Ocimum sanctum (Tulsi) leaves are used for cough, cold and fever,
(ii) Mentha spicata (Podina) leaves, used for preparing ‘Chutney’, help in indigestion and
rheumatism, (iii) Thymus vulgaris leaves are effective against hookworms, and (iv) Mentha
longifolia leaves are used in rheumatic pain and some heart diseases.
• Camphor is obtained from the oil of Ocimum kilimandscharicum.
• Root tubers of some species of Coleus and Ocimum are edible.

Labiatae has been placed under order Lamiales in majority of the systems of classification.
Briquet in Engler and Prantl’s Pflanzenfamilien divided Labiatae into 8 subfamilies (Ajugoideae,
Prostantheroideae, Prasioideae, Scutellarioideae, Lavanduloideae, Lamioideae, Ocimoideae and
Catopherioideae).
422 Plant Taxonomy
Labiatae is closely allied to Verbenaceae as well as to Boraginaceae. In some minor characters

it also shows affinities with Scrophulariaceae. Hutchinson opined that order Lamiales, to which
Labiatae belongs, is the most highly evolved order among Dicotyledoneae.

Ocimum basilicum (Sweet Basil or Ban Tulsi, Fig. 21.98)
Habit: Cultivated aromatic herb. Stem: Herbaceous, erect, hairy, quadrangular. Leaf: Simple,
opposite decussate, petiolate, exstipulate, ovate, serrate, hairy, unicostate reticulate. Inflorescence:
Verticillaster. Flower: Bracteate, pedicellate, complete, hermaphrodite, hypogynous, zygomorphic,
pentamerous. Calyx: 5 sepals, fused, 1/4 bilabiate, valvate. Corolla: 5 united petals, 4/1 bilabiate,
valvate. Androecium: 4 stamens, epipetalous, didynamous, posterior stamen absent; anthers dithecous,
dorsifixed, introrse. Gynoecium: Same as in general description of the family. Fruit: Schizocarpic
(carcerulus), consisting of 4 nutlets.
Floral Formula: Br, , , K(1/4), C (4/1) A2+2, G (2).
stigma stigma anther lobe
stamen filament
flower
petal
larger
petal style
ovary
sepal
disc sepal
pedicel ovule pedicel
A Flower L.S. Flower

inflorescence
stigma
leaf
style
stem
disc
ovary Flowering Branch

Gynoecium
Floral Diagram
Fig. 21.98 Ocimum basilicum L.

21.85 VERBENACEAE VERBENA FAMILY

Gamopetalae, Bicarpellatae, Lamiales.

Herbs, shrubs, or small trees; stem usually quadrangular; leaves usually opposite; flowers zygomor-
phic, pentamerous; ovary not lobed; style terminal.

Clerodendrum, Duranta, Lantana, Tectona, Verbena, Vitex.

A family of about 99 genera and 3151 species (Radford, 1986), distributed chiefly in tropics and
subtropics. About 22 genera and over 30 species have been reported from India. Some of the genera
along with their number of approximately reported species and common names of some of them are
Clerodendrum (400, glory-bower), Verbena (250, vervain), Vitex (250, chaste tree), Lippia (220, frog-
fruit), Lantana (150), Callicarpa (140, beauty-berry), Stachytarpheta (100) and Tectona (3, teak).

General Habit Herbs, shrubs (Lantana) or trees (Tectona); often lianas (Clerodendrum, Vitex), some-
times of a xerophytic nature (several species of Verbena); a few are mangrove shrubs (Avicennia).
Stem Herbaceous or woody, twining or sprawling in climbers; often quadrangular.
Leaf Usually simple, sometimes pinnately (Peronema) or palmately (Vitex) compound; usually
opposite, rarely alternate or whorled; exstipulate; highly reduced in xerophytic species (Verbena).
Inflorescence Racemose raceme (Duranta), or umbel (Lantana) or a spike; often with an involucre
of coloured bracts; or consisting of dichotomous cymes, or panicled cymes (Tectona).
Flower Bracteate, often bracteolate (Duranta, Lantana), complete, hermaphrodite, zygomorphic
(rarely actinomorphic, e.g. Physopsis), pentamerous, hypogynous.
Calyx Usually 5 united sepals, persistent; usually lobed or toothed; valvate; rarely the sepals are
4, 6 or 8; sometimes coloured.
Corolla Usually 5 united petals, or as many petals as sepals; often bilipped with a narrow tube;
rarely campanulate; typically lobed with the lobes generally unequal, usually imbricate.
Androecium Usually 4 stamens, didynamous, epipetalous, alternate with the corolla lobes; anthers
dithecous, introrse; longitudinally dehiscent; posterior stamen is generally missing or present in the
form of staminode; only 2 fertile stamens and 3 staminodes are present in Oxera; in Tectona all
the 5 stamens are fertile.
424 Plant Taxonomy
B C
Fig. 21.99A–C Verbenaceae: Floral diagrams–A: Duranta plumiri; B: Clerodendrum indicum, and
C: Verbena.
Gynoecium Usually bicarpellary, syncarpous; ovary superior; originally bilocular but divided into 4
locules by the formation of false septum in each locule; 2 ovules in each carpel i.e. 1 ovule in each
locule after septation; sometimes 2 ovules in each locule (Fig. 21.99A); axile placentation; ovules
anatropous to orthotropous; style terminal and only rarely sunk in between the lobes of the ovary;
stigma lobes usually as many as carpels; rarely carpels are 4 (Duranta) or 5 (Geunsia); ovary is
unilocular with free-central placentation in Avicennia; in Clerodendrum and Verbena the ovary is
unilocular with parietal placentation when young but on maturity the placentae unite at the centre
and the placentation becomes axile (Fig. 21.99B, C).
Fruit and Seeds Fruit is generally a drupe, and rarely a capsule, berry, or schizocarp (nutlets). Seeds
non-endospermic with straight embryo.
Pollination and Dispersal Pollination is mainly by butterflies and bees, and seeds are dispersed
mainly by birds or animals.
General Floral Formula Br, Brl, , , K(5), C (5), A2+2, G (2).

• Timber-yielding Plants: Teak (Tectona grandis), one of the most famous timber-yielding
plants of the world, belongs to Verbenaceae. It is used for high quality furnitures, heavy
constructions, ship building, musical instruments and several other similar purposes. Some
other timber-yielding plants of Verbenaceae are Gmelina arborea, Premna benghalensis and
Vitex altissima.
• Common garden ornamentals include Aloysia triphylla, Caryopteris incana, Clerodendrum
bungei, C. fragrans, C. interne, C. siphonanthus, C. splendens, C. thomsoniae, C. trichoto-
mum, Duranta repens syn. D. plumieri (golden dewdrop), Holmskioldia sanguinea, Lantana
camara, Petrea volubilis (Queen’s wreath), and several species of Verbena (Vervain) such
as V. bipinnatifida and V. hybrida.
• Verbena oil, used in perfumery, is obtained from the leaves of Aloysia triphylla.

Verbenaceae was placed under Lamiales by majority of the workers including Bentham and Hooker,
Takhtajan, Cronquist, and Thorne. However, Hutchinson placed it under order Verbenales. Briquet
divided Verbenaceae into 7 tribes (Stilbeae, Verbeneae, Chiloantheae, Viliceae, Caryopterieae,
Symphoremeae and Avicenneae).
Verbenaceae is closely allied to Labiatae, and to some extent to Boraginaceae and Scrophulariaceae.
However, Hutchinson opined that Verbenaceae and Labiatae are not at all related.

Lantana camara Linn. (Fig. 21.100)
Habit: A small shrub. Leaf: Simple, exstipulate, opposite decussate, petiolate, ovate, crenate or ser-
rate, acute, unicostate reticulate. Inflorescence: Umbel or a compound spike with flat top. Flower:
Bracteate, bracteolate, hermaphrodite, zygomorphic, pentamerous, hypogynous. Calyx: 5 sepals,
united, valvate. Corolla: 5 petals, united, quincuncial; corolla bilipped (4/1) in which 1 anterior petal
is large and pointed; white, yellow, red or pale-purple. Androecium: 4 stamens, didynamous, epipeta-
lous; posterior stamen is generally missing; dithecous, basifixed, introrse. Gynoecium: Bicarpellary,
syncarpous, superior, bilocular, 1 ovule in each locule, axile placentation; style simple; stigma knob-
like. Fruit: Drupe.
Floral Formula: Br, Brl, , , K(5), C (4/1), A2+2, G (2).
21.86 PLANTAGINACEAE PLANTAGO FAMILY

Gamopetalae, Bicarpellatae, Ordines Anomali.
Or
Metachlamydeae (Sympetalae), Plantaginales (Engler and Prantl).
426 Plant Taxonomy
stigma anther
petal lobe
stigma connective
style
stamen
style filament
ovary
ovule
A Stamen
sepal flower
Gynoecium
ovary
pedicel
bract
L.S. Flower bract
leaf
stem
Floral Diagram
Flowering Branch
Fig. 21.100 Lantana camara Roxb.
Or
Dicotyledones, Herbaceae, Plantaginales (Hutchinson).
Or
Magnoliopsida, Asteridae, Plantaginales (Cronquist, 1981).

Herbs; leaves in a basal rosette with apparent parallel venation and often sheathing at base; spicate
or capitate inflorescence on stout or wiry scapes; flowers usually tetramerous; petals membranous;
stamens often exerted.

Plantago.

A family of only 3 genera (Plantago, Littorella and Bougueria) and over 250 species, Plantaginaceae
are cosmopolitan in their distribution. Plantago (Plantain) with more than 200 species is the largest
genus. Bougueria is a monotypic Andean genus.

General Habit Annual or perennial herbs or rarely well-branched subshrubs.
Leaves Basal, simple, alternate or rarely opposite; often sheathing at the base, narrow, parallel-
veined with no distinction into stalk and blade; exstipulate; in Plantago lanceolata the leaves form
a basal rosette (Fig. 21.101), often lanceolate to ovate, entire, 3–5 nerved and sessile.
spike of
flowers stigma
stamens (4)
style petal
scape
leaves
sepals
bract E
A Mature Single Flower
spike of
flowers
Flowering Plants scape
G
A Young Fruit
B
stigma Top of Furrowed Scape
floral buds
hairs inflorescence
axis
corolla
tube
hairy
sepals
bract D C ovary F
L.S. Top of Young
A Single Flower L.S. Superior Ovary
(Young) Inflorescence
Fig. 21.101 Plantago lanceolata L.

428 Plant Taxonomy
Inflorescence Scapose, capitate or spicate present on stout or wiry scapes: scapes usually twice as
long as leaves (Plantago, Fig. 21.101).
Flowers Usually small, inconspicuous, bracteate, bisexual, actinomorphic, hypogynous,
tetramerous
Calyx Sepals 4, gamosepalous, imbricate; sometimes deeply-divided and membranous.
Corolla Petals 4, gamopetalous, imbricate and membranous.
Androecium Stamens 4, rarely 1–2, epipetalous, alternipetalous; anthers dithecous, versatile, large;
dehiscing generally longitudinally.
Gynoecium Bicarpellary, syncarpous, ovary superior, 1 to 4 locules with 1 or more ovules in each
locule on basal or axile placentation; usually only 2 locules; style simple, filiform and bifid.
Fruit A capsule or bony nutlet (e.g. Littorella) surrounded by persistent calyx.
Seeds With fleshy endosperm and small straight embryo.

Members of Plantaginaceae are not of much economic importance except of the following:
• Several species of Plantago are troublesome garden weeds, specially in lawns.
• Seeds of Plantago psyllium are used as a laxative. The mucilaginous seed coats of P. indica
and P. ovata are also used for this purpose.
• Leaves of some species of Plantago have occasionally been used as food.

Plantaginaceae is an anomalous taxon and hence of doubtful relationship. Due to this Bentham and
Hooker treated this family under Ordines Anomali. Bessey included it in Primulales as allied to and
originating from Plumbaginaceae. Hallier discussed it under Tubiflorae as related to Scrophulariaceae.
Hickey and King (1988) also regarded Plantaginaceae as derived from a pentamerous type of flower
as that of Veronica of Scrophulariaceae, and due to this the two families are considered as being
allied. Both Lawrence and Rendle retained it under order Plantaginales as also by Hutchinson
and Cronquist. Some taxonomists believe that Plantaginaceae represents “an offshoot from the
Polemoniales or perhaps from the Primulales” (Porter 1959).
21.87 MONOCHLAMYDEAE
Instead of free or distinct petals of Polypetalae and completely or partially fused petals of
Gamopetalae, members of Monochlamydeae have no petals. Monochlamydeae is directly divided
into following 8 series with no cohorts by Bentham and Hooker:
Series 1. Curvembreae, characterised by coiled embryo and generally one ovule. It includes 7 orders
(=families), viz. Chenopodiaceae1, Polygonaceae1, Nyctaginaceae, Amaranthaceae1, Illeceraceae,
Phytolacaceae and Batideae.
1
Discussed in this text.
Series 2. Multiovulatae-aquaticae, characterised by their aquatic or immersed habit and presence of

syncarpous pistil and many ovules. It includes Podostemaceae.
Series 3. Multiovulate-terrestris, characterised by their terrestrial habit and presence of syncarpous pistil
and many ovules. It includes families such as Nepenthaceae, Aristolochiaceae1, and Cytinaceae.
Series 4. Microembryeae, characterised by the presence of very minute embryo in copious endosperm,
apocarpous or syncarpous condition, and usually single ovule. It includes Piperaceae1, Chloranthaceae,
Myristicaceae and Monimiaceae.
Series 5. Daphnales, characterised usually by monocarpellary gynoecium, ovary with one carpel and
one ovule, bisexual flowers and mostly woody plant. It includes Proteaceae, Lauraceae, Thymeleaceae,
Elaeagnaceae and Penaeaceae.
Series 6. Achlamydosporeae, characterised by usually inferior ovary, single locule with 1–3 ovules;
seeds without a seed coat; often parasitic habit. It includes Loranthaceae1, Santalaceae and
Balanophoraceae.
Series 7. Unisexuales, characterised by unisexual flowers. It includes 9 families, viz. Euphorbiaceae1,
Urticaceae1, Moraceae1, Plantanaceae, Leitneriaceae, Juglandaceae, Balanopsidaceae, Myricaceae and
Casuarinaceae1.
Series 8. Ordines anomali, includes families of uncertain relationship. These bear generally unisexual
flowers but their plants are not closely related to others. It includes families like Ceratophyllaceae,
Salicaceae1, Lacistemaceae and Empetraceae.
21.88 CHENOPODIACEAE (GOOSEFOOT FAMILY)

Monochlamydeae or Apetalae, Curvembryeae.

Halophytic, xerophytic or succulent, annual or perennial herbs or shrubs; leaves and stem mealy to
touch; flowers monochlamydous, small and greenish; placentation basal; fruit urticle.

Basella, Beta, Chenopodium, Kochia, Spinacia and Suaeda.

A family of about 102 genera and over 1400 species, Chenopodiaceae are cosmopolitan in distribu-
tion, found chiefly in saline and xeric habitats. About 20 genera and over 55 species represent the
family in India. Some of the genera along with their common names and approximate number of
their worldover reported species are Atriplex (200, saltbush), Salsola (150, glassworts), Chenopodium
(150, goosefoot), Kochia (90, summer-cypress), Salicornia (35, glassworts), Beta (6, sugarbeet or
Chukander), and Spinacia (3, spinach or Palak).
1
Discussed in this text.
430 Plant Taxonomy

General Habit Predominantly halophytic (Atriplex, Salsola kali) or xerophytic; annual or peren-
nial herbs or shrubs, only rarely small trees (Haloxylon ammodendron), or sometimes succulent
(Salicornia); mostly adapted to grow in salty or alkaline soil; plant parts covered by hairs.
Roots Deep-rooted; become thick, fleshy and store large amount of sugar in some (e.g. Beta
vulgaris).
Stem Jointed, or sometimes becomes fleshy and leafless in succulents (e.g. Salicornia); shows
abnormal secondary growth by forming interxylary phloem.
Leaf Simple, alternate, exstipulate; rarely opposite (Haloxylon, Salicornia); fleshy in some (Suaeda)
or reduced to scales; often covered with hairs which provide a characteristic mealy appearance to
the plants.
Inflorescence Usually cymose clusters (Chenopodium) arranged in terminal or axillary dense spikes;
often dense, dichasial or unilateral cymes.
Flower Bracteate, complete, bisexual, or sometimes unisexual (Grayia), and the species may be
monoecious (Sarcobatus) or dioecious; actinomorphic, hypogynous but rarely perigynous (Beta);
monochlamydous; very small; greenish; Atriplex has male, female and sometimes also bisexual
flowers.
Perianth Usually 5 tepals, sometimes varying between 2 to 5; free or connate at the base; persistent
in the fruit; quincuncial or imbricate; mostly fleshy, small, green and resemble sepals; tepals differ
in shape and size (Alexandra, Corispermum); petals absent.
Androecium Usually 5 or as many stamens as tepals, polyandrous, opposite the tepals (Fig. 21.102);
anthers bent inwards in bud; filaments usually free or subulate; anthers dithecous, introrse, longi-
tudinally dehiscent; only 1 or rarely 2 stamens in Salicornia; anthers open by terminal pores in
Basella.
Gynoecium Usually bi-or tricarpellary, syncarpous, superior or rarely half-inferior (Beta); unilocular,
with one campylotropous ovule, basal placentation; styles 1 to 3; stigmas 2–3 or rarely more.
Fruit and Seeds Fruit usually a small, round utricle, nut or achene. Seed single in each fruit, lens- or
kidney-shaped; with curved or spirally coiled embryo, usually surrounding the endosperm; sometimes
non-endospermic
General Floral Formula Br, ≈, , P5, A5, G (2 or 3).

• Sugar: Tuberous napiform roots of Beta vulgaris (sugarbeet or Chukander) contain about
20% sugar, and are the chief source of sugar in several temperate countries.
• Edible Plants: (i) Green leaves of Atriplex hortensis (Orache), Chenopodium album
(Bathua), C. bonus-henricus (Good King Henry), Beta benghalensis (Gobru Palak)
and Spinacia oleracea (Spinach or Palak) are used as famous vegetables. (ii) Seeds of
stigma anther lobe

stamen
stigma
style style connective
tepal ovary
ovule filament
ovary
Gynoecium
pedicel A Stamen
L.S. Flower
inflorescence
leaf
stem
Fig. 21.102 Chenopodium album L.
Chenopodium album and C. quina are ground into flour and eaten. (iii) Roots of Beta
vulgaris are eaten as ‘salad’. (iv) Seeds of Chenopodium quina are boiled and eaten with
rice.
• Fodder for cattles is obtained from Atriplex hortensis, Chenopodium album, Kochia indica
and Salsola foetida.
• Ornamental plants of the family are Kochia scoparia (Burning bush), Suaeda maritima,
Chenopodium amaranticolor and Atriplex hortensis.
• Chenopodium ambrosioides is used against hookworms.

Treated under series Curvembryeae by Bentham and Hooker, Chenopodiaceae was placed under
Centrospermae by Engler and Prantl (1931), under Chenopodiales by Hutchinson (1959) and
432 Plant Taxonomy
Thorne (1983), and under Caryophyllales by Takhtajan (1969) and Cronquist (1981). Bentham and
Hooker divided Chenopodiaceae into two subfamilies (Chenopodieae and Baselleae) while Ulbrich
(1934) in Engler and Prantl’s Pflanzenfamilien divided it into 8 sub-families (Polycnemoideae,
Betoideae, Chenopodioideae, Corispermoideae, Salicornioideae, Sarcobatoideae, Suaedoideae and
Salsoloideae).
Chenopodiaceae is closely allied to Amaranthaceae, from which it differs in not possessing mem-
branous perianth and connate stamens.
21.89 AMARANTHACEAE AMARANTHUS OR PIGWEED FAMILY


Mostly herbs; dense or congested inflorescence; flowers small with dry scarious bracts; stamens
connate, at least at the base; fruit utricle or capsule.

Achyranthes, Alternanthera, Amaranthus, Celosia, Digera, Gomphrena and Pupalia.

A family of about 65 genera and 900 species, Amaranthaceae are mostly distributed in tropical but
also in temperate regions. About 18 genera and over 50 species have been reported from India. Some
of the larger genera with the number of their approximately reported species and common names are
Alternanthera (200, alligator weed), Gomphrena (100, globe amaranth), Iresine (80, gizzard plant),
Amaranthus (60, pigweed), and Celosia (60, Celosia).

General Habit Mostly annual or perennial weedy herbs, rarely woody shrubs or climbing shrub
(Deeringia amaranthoides).
Stem Anomalous secondary growth occurs by the formation of collateral vascular bundles from cam-
bium in the pericycle. In the scattered vascular bundles, the fascicular cambium is functionless.
Leaf Simple, alternate (Amaranthus) or opposite decussate (Achyranthes, Fig. 21.103), exstipulate,
usually entire or nearly so, often thickly covered with hairs.
Inflorescence Dense or congested, axillary or terminal spikes (Amaranthus), heads, racemes; or
flowers solitary; often spiny because of the firm spiny tips on the inflorescence bracts; usually
the basic unit is either a 3-flowered dichasium unit (Celosia) or 7 to 11-flowered dichasium units
(Pupalia).
Flower Bracteate, bracts scaly, membranous, dry or scarious; bracteolate, bracteoles 2, dry, membra-
nous, large and lateral; usually bisexual, or when unisexual (Amaranthus, Fig. 21.104B, C) the species
are mostly dioecious or polygamodioecious; actinomorphic, pentamerous, hypogynous, incomplete,
apetalous, very small.
stigma
style
stamen
stamen
staminode ovary
ovule
tepal
Stamens and Staminodes

L.S. Flower
inflorescence
bracteole
A Flower
leaf
stem
Floral Diagram
Flowering Branch
Fig. 21.103 Achyranthes aspera L.
Perianth Uniseriate, consisting of 5, free or basally connate tepals; green or sepalloid, dry or
membranous; imbricate; only 3 tepals in Amaranthus viridis, but 5 free tepals in A. spinosus
(Fig. 21.104).
Androecium Usually 5 stamens present opposite the tepals; often monadelphous (Celosia,
Achyranthes); separate in Amaranthus, often with equal number of staminodes alternating with
the anthers (Celosia, Achyranthes); staminodes are fimbriated in Achyranthes; filaments united at
the base into a short tube, but in Amaranthus the filaments are free; anthers dithecous, dorsifixed
(Achyranthes) or versatile (Amaranthus); introrse; longitudinally dehiscent; only 1 or 2 stamens in
Nothosaerva.
Gynoecium Bi- to tricarpellary, syncarpous, superior, unilocular, with a single pendulous or campy-
lotropous ovule in the locule, basal placentation; rarely several ovules (Celosia, Fig. 21.104A) are
present; styles 1 to 3; stigma 1 to 3, capitate.
434 Plant Taxonomy
C
B
Fig. 21.104A–C Amaranthaceae: Floral diagrams—A: Celosia; B and C: Amaranthus spinosus, male
and female flower, respectively.
Fruit and Seeds Fruit usually a utricle (Achyranthes) or nutlet (Digera), or a capsule (Celosia), or
rarely a berry or drupe. Seeds with curved embryo and mealy endosperm.
Pollination and Dispersal Pollination is entomophilous because numerous small flowers are aggre-
gated in dense or congested inflorescence. Dispersal takes place either by wind or by animals.
General Floral Formula Br, Brl, ≈, , P5, A(5+5 Staminodes), G (2 or 3).

• Ornamental plants of the family known for their multicoloured foliage and beautiful inflores-
cence are Amaranthus caudatus (Foxtail), A. tricolor, A. salicifolius, Celosia cristata (cocks
comb; white, purple or yellowish-red panicles), Deeringia amaranthoides, Gomphrena glo-
bosa (globe amaranth), Iresine herbtsii (purplish-red foliage) and I. lindenii (red foliage).
• Edible plants of the family, of which the leaves and young shoots are used as vegetable, are
Amaranthus blitum (Amaranth or Chaulai), A. caudatus, A. hybridus, A. spinosus (Kantili
Chaulai), A. tricolor (Bari Chaulai), A. viridis, Celosia argentea and Digera muricata
(Lehsua). Grains of Amaranthus caudatus are roasted and eaten as ‘Ramdana’.
• Inflorescence paste of Achyranthes aspera (Latjeera or Chirchita) is used as an antidote
against snake and scorpion bites.
• Widespread troublesome weeds of this family belong to several genera including Acnidia,
Amaranthus, Gomphrena and Iresine.

Amaranthaceae was differently placed under series Curvembreae (Bentham and Hooker), Centrospermae
(Engler and Prantl), Chenopodiales (Hutchinson, 1959; Thorne, 1983) and Caryophyllales (Takhtajan,
1969; Cronquist, 1981). Bentham and Hooker divided Amaranthaceae into three tribes viz. Celosieae,
Amaranteae and Gomphreneae.
The family is closely related to Chenopodiaceae, from which it differs in having dry and mem-
branous perianth, quite conspicuous and scarious bracts and bracteoles, and usually connate stamens.
Hutchinson opined that Amaranthaceae evolved from Caryophyllaceous ancestors.
21.90 POLYGONACEAE BUCKWHEAT OR SMARTWEED FAMILY


Herbs with swollen nodes; ochreate stipule present; tepals petalloid and often biseriate; petals absent;
fruit usually lens-shaped or triangular.

Antigonon, Coccoloba, Fagopyrum, Polygonum, Rheum, Rumex.

Represented by about 40 genera and 1000 species, Polygonaceae are distributed chiefly in north
temperate regions, and a few also in tropical, arctic and Southern Hemisphere. About 10 genera
and over 100 species have been reported from India. Some of the larger genera along with their
number of approximately reported species are Polygonum (300, smartweed), Eriogonum (250, false
buckwheat), Rumex (200, dock), Coccoloba (150, seagape), Rheum (50, rhubarb), Chorizanthe (50),
Fagopyrum (15) and Muehlenbeckia (15).
Polygonum orientale, an ornamental plant, is commonly known as “kiss-me-over-the-garden-gate”
(Jones and Luchsinger, 1987).

General Habit Mostly herbs, some shrubs (Polygonum hydropiper) or climbers (Antigonon leptopus),
and only rarely trees (Triplaris, Coccoloba uvifera).
436 Plant Taxonomy
Stem Often with swollen nodes; sometimes geniculate i.e. bent like a knee; phylloclades in some
species (Muehlenbeckia platyclada).
Leaf Usually simple, alternate, stipulate; stipules ochreate i.e. stipules united into a sheath (ochrea)
clasping the stem above the leaf base, a diagnostic feature of Polygonaceae; rarely the leaves are
opposite or whorled; usually entire but lobed in Rumex acetosella; leaves radical in R. hastatus;
rarely exstipulate (Koenigia islandica).
Inflorescence Primarily racemose, but partial inflorescences usually cymose; clusters of pani-
cled raceme (Rumex); or racemose forming a terminal panicle (Polygonum); or cymose umbels
(Eriogonum).
Flower Bracteate, pedicellate; usually bisexual but rarely unisexual; when unisexual, the species may
be monoecious or dioecious; actinomorphic; small and crowded on the inflorescence axis; usually
trimerous but rarely dimerous (Oxyria digyna), hypogynous.
Perianth 3 to 6, free or basally connate tepals, often in two whorls (Rumex); monochlamydous; often
petalloid; in acyclic flowers (Polygonum glabrum) 5 tepals are present; imbricate; often persistent,
enlarged and becoming membranous in fruit; petals absent.
Androecium 6 to 9 stamens, in two series (rarely less or more), free or united at the base; anthers
2-locular; dehiscence longitudinal; in several genera the outer stamens are introrse while the inner
stamens are extrorse; only 5 to 8 stamens (5 introrse and 2 extrorse) in Polygonum glabrum
(Fig. 21.105).
Gynoecium Usually tricarpellary, rarely bicarpellary (Oxyria) or tetracarpellary; syncarpous, supe-
rior, unilocular, one orthotropous ovule, basal placentation; style 1; stigmas 2 to 4, fringed in Rumex;
usually with an annual, nectar-secreting, hypogynous disc is present.
Fruit and Seeds Fruit a lens-shaped, angular, usually triangular achene or nut (Polygonum, Rumex)
with a persistent perianth forming a membranous wing. Seeds with a curved embryo and plenty of
endosperm.
Pollination It is with the help of wind in Rumex and some species of Polygonum, or by insects in
Rheum and several other species of Polygonum.
General Floral Formula Br, ≈, , P3+3, A6–9, G (3).

Except of some medicinal and ornamental value, the family is not of much importance. Some of
its economic aspects are undermentioned:
• Common ornamental plants are Antigonon leptopus (climber with bright pink or white flow-
ers), Muehlenbeckia platyclada syn. Coccoloba platyclada, Polygonum alpinum, P. glabrum
(pink-flowers), P. orientale, P. paniculatum and P. blebejum (pink flowers).
• Some of the medicinal plants of family include (i) Polygonum aviculare, used against diabetes
and rheumatism, (ii) Rheum emodi and R. officinale roots and rhizome yield the drug ‘rhu-
barb’ used as a laxative, purgative and stomach tonic, (iii) Rumex acetosella leaves are said
stigma stamen stigma
style
stamen
tepal
style tepal
ovule
pedicel ovary
bract
pedicel
ovary A Flower L.S. Flower
Gynoecium inflorescence
leaf
ochreate stipule
stem
Fig. 21.105 Polygonum glabrum Willd.
to be effective in tumours and cancer, and (iv) Rumex acetosa, used in bronchial diseases
and also as a blood purifier; (v) Polygonum glabrum seeds are used in leucorrhoea.
• Leaves of Oxyria digyna, Rumex acetosella and R. acetosa (sorrel) are eaten as vegetable
or salad.
• Fagopyrum esculentum (buckwheat or ‘Kuttu’) seeds are edible because they are rich source
of protein. Its flowers are very rich in honey. F. tataricum (duckwheat) is also used for the
similar purposes.
• Rumex hastatus leaves are used as condiment.
• Polygonum tinctorium is the source of a blue dye.
438 Plant Taxonomy

In most of the systems of classification (Hutchinson, 1959; Takhtajan, 1969; Cronquist, 1981; Thorne,
1983), the family Polygonaceae is placed under order Polygonales. Bentham and Hooker placed
it under series Curvembryeae of Monochlamydeae. The family is divided into 3 subfamilies, (i)
Rumicoideae (flowers cyclic and endosperm not ruminate), (ii) Polygonoideae (flowers acyclic and
endosperm not ruminate), and (iii) Coccoloboideae (flowers acyclic and endosperm ruminate).
Polygonaceae is allied to families such as Amaranthaceae, Chenopodiaceae and Caryophyllaceae.
According to Hutchinson Polygonaceae and families of Caryophyllales were derived from a common
ancestor which originated from Ranales.
21.91 ARISTOLOCHIACEAE BIRTHWORT FAMILY

Monochlamydeae, Multiovulatae Terrestris.

Plants terrestrial herbs or shrubs (mainly lianas); leaves alternate, often cordate; several stamens
(6–36) showing adonation to style; 6-loculed or more or less inferior ovary; flowers lacking corolla;
perianth usually an enlarged and a petalloid gamosepalous calyx often of trumpet-like or bell-like
shapes; ovules many.

Aristolochia, Asarum.

A family of 8–10 genera and about 600 species distributed in tropical and temperate America, Asia,
Europe and Africa; absent in Australia; Aristolochia (500) and Asarum (70) are the two larger genera
found mostly in tropical and temperate regions.

General Habit Herbs or shrubs, the latter mostly twining lianas.
Stem Softly woody with broad medullary rays.
Leaf Usually simple, alternate, petiolate, often ovate-cordate and palmately-veined; entire;
exstipulate.
Inflorescence Flowers in axillary clusters (Aristolochia, Fig. 21.106), solitary or arranged in racemes
or cymes.
Flower Bisexual, actinomorphic (Asarum) or zygomorphic (Aristolochia), epigynous.
Perianth Usually a petaloid gamosepalous calyx, variously 3-lobed or unilateral, often bizarrely
coloured; corolla usually absent but sometimes an inner whorl of 3 minute teeth (or vestigial corolla?)
hairs
pointing-downward
utricle
surface of
stigma
anther
gynostemium
inferior ovary
leaf
C
L.S. Utricle
stem curved perianth

tube
flowers utricle
B
A A Flower With
A Flowering Branch Inferior Ovary
gynostemium
leaf
inferior
ovary
fruit E
T.S. Ovary
F D
A Fruit Inferior Ovary With
Gynostemium
Fig. 21.106 Aristolochia clematitis L.

440 Plant Taxonomy
present; in Aristolochia clematitis (Fig. 21.106) three united perianth lobes form a yellow tube with
a brownish limb.
Androecium Stamens 6–36, free, or united with the style to form staminal column (gynostemium);
filaments short and thick; anthers free or adnate to style, dithecous, dehiscing longitudinally.
Gynoecium Compound pistil of 4–6 united carpels; with mostly 4–6 locules with numerous anat-
ropous, horizontal or pendulous ovules in each locule; axile or parietal placentation; ovary inferior,
rarely half-inferior; style 1, short and stout; with 4–6 stigmas.
Fruit and Seed Fruit a many-seeded or septicidal capsule, often dehiscing basally i.e. parachute-like.
Seed with small embryo, rich in endosperm.
General Floral Formula EBr, ≈ or , , P(3), A6–36, G (4–6).

Not being of much economic importance, many species of two genera (Asarum and Aristolochia)
of the family have some horticultural value and cultivated as ornamentals. Aristolochia macro-
phylla is often grown as ornamental vine and called Dutchman’s pipe because of its bent perianth
tube and brownish flowers. A serpentina (Virginia snakeroot) has been used medicinally and
serve as a cure for snake-bites. Asarum (wild ginger) also includes several species of horticultural
interest. Roots of Aristolochia indica are used by snake-charmers for catching snakes. Some
other species of ornamental value include A. clematitis (Fig. 21.106), A. gigas, A. grandiflora, A.
ornithocephala and Asarum europaecum.

Aristolochiaceae has been included under series Multiovulatae Terrestris of Monochlamydeae
by Bentham and Hooker. However, it has been treated under order Aristolochiales of subclass
Archichlamydeae by Engler and Prantl and under the same order (Aristolochiales) of class
Magnoliopsida by Takhtajan (1980) and Cronquist (1981). Hickey and King (1988) have mentioned
that inspite of many attempts to find a suitable position in the system for the Aristolochiaceae, “it
has proved a difficult family to place, and there is still a divergence of opinion on this matter. It is,
however, probable that it is connected with the Annonaceae, a family in the Magnoliales, through
the genera Thottea and Apama”. Engler divided the Aristolochiaceae into three tribes, namely
Sarumeae (flowers regular and solitary), Bragantieae (flowers regular but in cymes or racemes) and
Aristolochieae (flowers irregular). Hutchinson (1948) placed Aristolochiaceae as a terminal derivative
from the Ranales via herbaceous members of the Berberidaceae.
21.92 PIPERACEAE PEPPER FAMILY

Monochlamydeae or Apetalae, Microembryeae

Succulent herbs a shrubs, rarely trees; petals absent; stamens 1–10; ovary unilocular; embryo very
minute.

Piper, Peperomia.

According to some authors, Piperaceae is composed of 12 genera and about 1400 species inhabiting
mostly rainforests. Hickey and King (1988), however, mention that Piperaceae is a family of only 8
genera and 3100 species. Piper, distributed mainly in tropics, is represented by about 2000 species,
while Peperomia is distributed both in tropics and subtropics with its 1000 species. Fresh leaves of
Piper betle are chewed with betlenuts and used as a masticatory in most of the Asiatic countries;
Peperomia incana (Fig. 21.107) is often grown as a pot plant in greenhouses, and it roots readily
from stem cuttings.

General Habit Mostly slender (Peperomia pellucida) or stout (P. tetraphylla) herbs, found in waste
places; P. incana (Fig. 21.107) is a fleshy subshrub, reaching about 30 cm in height; often climbing;
more rarely small trees.
Stem Herbaceous, sometimes succulent, often climbing; distinct vascular strand scattered and not
arranged in concentric rings.
Leaf Alternate, rarely opposite or whorled; simple, petiolate, entire, sometimes fleshy; stipules,
if present, are lateral adnate; in Peperomia incana (Fig. 21.107) leaves are stiff, broadly cordate,
covered with hairs.
Inflorescence A dense spike or umbellate spikes; spikes of usually unisexual flowers in Piper.
Flower Minute, bracteate, usually bisexual, devoid of perianth; hypogynous, regular; in Peperomia
incana (Fig. 21.107) the flowers are minute and deeply sunk into the fleshy axis of the inflorescence,
posing difficulty even in seeing the floral parts.
Perianth Absent.
Androecium Stamens 1–10; anthers 2-celled (Piper), confluent into one in Peperomia; dehiscence
longitudinal.
Gynoecium Carpels 1–5, united; ovary superior, unilocular; basal orthotropous ovule, one in each
locule; stigmas 1–5.
Fruit A small drupe.
Seeds Very small, embryo minute; with abundant mealy perisperm.
Pollination Pollination process is little known in Piperaceae. According to Hickey and King (1988)
Piperaceae are considered to be closely related to the insect-pollinated Magnoliaceae showing ento-
mophily. The flowers are protogynous, which encourages cross-pollination. Due to glutinous nature
of pollen grains and scented flowers, at least Peperomia resediflora is insect-pollinated.
General Floral Formula Br, ≈, , P0, A1–10, G (1–5).
442 Plant Taxonomy
ovary
stigma with
tuft of papillae
stamen
spike floral bract
B inflorescence
axis
leaf
L.S. of Flower in Fleshy Axis
ovary
stamen
ovule stigmatic
paillae
inflorescence
axis C
T.S. Flower in Fleshy Axis
shoot
A stigma spike
Flowering Branch stamen

ovary
closely-set
flowers
stigma G
A Part of T.S. Inflorescence
ovary Showing A Single Flower
stamens apetalous
flowers
bract leaf
axis
shoot
E D
F
Apetalous Flower T.S. Inflorescence Axis Terminal Spicate
(Anterior View) Inflorescence
Fig. 21.107 Peperomia incana.


• Piper betle (Betel or Paan) is used as a common masticatory in many Asiatic countries. Its
fresh leaves are chewed with betlenuts.
• The whole drief fruits of Piper nigrum are known as “black pepper”. If the outer layer is
removed, the fruits are called “white pepper”.
• Dried unripe fruits of Piper cubeba are used for flavouring as the source of “cubebs”,
while the roots of P. methysticum are used for making an intoxicating Polynesian beverage
“Kava”. P. caninum yields a type of volatile oil called “Kababchini”. Wood of P. chaba
forms a pungent condiment. “Long pepper” is obtained from P. longum and P. retrofactum.
In Mexico, walking sticks are made from the stems of P. tiliaefolium.
• Many species of Peperomia and a few of Piper and Macropiper are grown for their orna-
mental value. Peperomia species are cultivated as house plants for their variegated foliage.

Bentham and Hooker included Piperaceae under series Microembreae of Monochlamydeae or
Apetalae while Engler and Prantl discussed it under order Piperales of subclass Archichlamydeae of
class Dicotyledoneae. Hutchinson (1973) treated Piperales under division Herbaceae of Dicotyledons.
The minute-flowered, spicate inflorescence of several members of Piperaceae bring it close to
Betulaceae and Corylaceae. Because of the entomophilous nature, however, Piperaceae are close to
Magnoliaceae.
21.93 LORANTHACEAE MISTLETOE FAMILY

Monochlamydeae or Apetalae, Achlamydosporae.

Usually aerial parasitic habit; cup-shaped receptacle; inferior ovary; ovules not differentiated from
the placenta.

Dendrophthoe, Loranthus, Viscum.

Family is represented by about 36 genera and 1300 species and distributed in both tropical and
temperate regions. About 10 genera and over 52 species have been reported from India. Some of
the largely represented genera along with the number of their approximately reported species are
Loranthus (600), Phoradendron (190), Struthanthus (75), Viscum (70), Dendrophthora (55) and
Psittacanthus (50).
444 Plant Taxonomy

General Habit Mostly semiparasitic herbs or small shrubs attached to their hosts by sucking haus-
toria; rarely erect, terrestrial trees (e.g. Nuytsia).
Roots Modified adventitious roots present in the form of haustoria; Nuytsia is rooted in the earth;
in Viscum the root often branches within the host tissue; haustoria penetrate the bark and sapwood
of the host.
Stem Usually dichotomously branched, often with swollen nodes.
Leaf Usually opposite or whorled, rarely alternate; exstipulate; sometimes reduced to scales; usually
evergreen, thick, leathery, entire and persistent.
Inflorescence Usually cymose, or a panicle, raceme, spike; or flowers solitary with cup-shaped
receptacle.
Flower Usually bracteate, bracteolate; bisexual (Loranthus) or unisexual (Viscum, Fig. 21.108);
usually actinomorphic, rarely slightly zygomorphic (Dendrophthoe); epigynous; flowers usually in
groups of 3 (or 2, by abortion of the central flower).
Perianth In 1 or 2 whorls of 4 to 6, free or united, large tepals developing from the margin of
a cup-shaped receptacle; tepals, if united, form a tube; sepaloid (Viscum) or petaloid (Loranthus);
calyculus, an outgrowth present in the form of a small rim or fringe below the perianth, develops
in several genera (Loranthus, Nuytsia, Struthanthus and Psittacanthus).
Androecium Stamens equal in number to perianth lobes, adnate to perianth lobes or at their base;
generally opposite the tepals; anthers 2-celled or transversely multiloculate; dehiscing either trans-
versely or longitudinally or even by terminal pores.
Gynoecium 3–5 carpels, syncarpous, inferior, unilocular, the ovules not differentiated from the
placenta; carpels usually sunk in or fused with the receptacle; basal placentation; style simple or
absent; stigma 1 and often sessile.
Fruit and Seeds: Fruit usually 1 to 3-seeded berry or drupe. Seeds usually surrounded by viscin,
a sticky substance; endospermic with straight embryo.
General Floral Formula: Br, Brl, ≈, or unisexual, P4–6 or (4–6), A4–6, G (3–5).

Plants of Loranthaceae are of no specific economic importance. Only Viscum album (Mistletoe,
Fig. 21.108) and some species of Phoradendron (e.g. P. flavescens) are of ornamental value and
used for decorative purposes, specially at Christmas time.

Most of the taxonomists place Loranthaceae under order Santalales (Lawrence, 1951; Takhtajan,
1969; Cronquist, 1981; Thorne, 1983). However, Bentham and Hooker discussed Loranthaceae under
series Achlamydosporae of Metachlamydeae. In Engler’s Syllabus der Pflanzenfamilien, Loranthaceae
is divided into 2 subfamilies viz. Loranthoideae (calyculus or 2 bracteoles present) and Viscoideae
(calyculus absent). Loranthaceae shows close affinities with Santalaceae.
leaf
male lateral
inflorescence shoot bud
young
inflorescence
pollen stamen stem

grains
fruits B
female
flowers
stem
A
C
Fig. 21.108 Viscum album L (A: Female plant; B: Male inflorescence; C: L.S. male flower; D: Female
inflorescence; E: L.S. female flower, and F: L.S. fruit).
21.94 EUPHORBIACEAE SPURGE FAMILY

Monochlamydeae or Apetalae, Unisexuales.

Herbs, shrubs, or small trees; milky latex present; flowers apetalous, unisexual; female flowers usu-
ally tricarpellary and ovary trilocular; fruit capsule or regma.
446 Plant Taxonomy

Croton, Euphorbia, Hevea, Jatropha, Manihot, Phyllanthus and Ricinus.

A family of about 300 genera and 7500 species (Jones and Luchsinger, 1987), Euphorbiaceae are
cosmopolitan in distribution. They are abundant in both tropical and temperate regions, and not
well-represented in arctic regions. Over 60 genera and about 350 species have been reported from
India. Some of the common genera along with the approximate number of their worldover reported
species as well as the common names of some of them are Euphorbia (2000, spurge or poinsettia),
Croton (750), Phyllanthus (600), Acalypha (450, chenille plant), Jatropha (175), Manihot (170),
Sapium (120), Tragia (100, nose-burn), Clutia (70), Manadenium (47), Ricinocarpos (16), Hevea
(12, Para-rubber) and Ricinus (1, castor bean).

General Habit Mostly shrubs (Ricinus communis, Jatropha gossypifolia) or trees (Emblica officina-
lis), or sometimes herbs (Phyllanthus nirurii), or climbers (Tragia involucrata); some are xerophytic
(many species of Euphorbia) or cactus-like or phylloclades; some are marshy (Coperonia); usually
the plants contain milky sap or latex.
Stem Herbaceous or woody; becomes cactus-like in several species of Euphorbia (e.g. E. neriifolia,
E. royleana, E. trigona); in E. xylophylloides stem becomes flattened and leaf-like (phylloclade).
Leaf Mostly simple (Euphorbia pulcherrima), alternate, stipulate; often reduced or deciduous as in
several xerophytic species of Euphorbia; sometimes palmately lobed (Ricinus, Jatropha), or deeply
so in Manihot; rarely opposite (Euphorbia hirta, Choriophyllum) or whorled (Mischodon); stipules
sometimes present in the form of hairs, glands, or thorns; venation is unicostate reticulate, or multi-
costate divergent; in Phyllanthus niruri leaves are arranged in 2 rows so that each branch resembles
with a compound leaf.
Inflorescence Highly variable from a raceme (Croton), a spike (Acalypha), a dichasium (Jatropha),
or even the flowers are solitary axillary (Phyllanthus asperulatus); but in majority of the cases, as
in Euphorbia, the inflorescence is a cyathium.
Each cyathium (Fig. 21.109) contains terminally a single, naked female flower, usually represented
by a tricarpellary gynoecium. The female flower is surrounded by a cup-like involucre formed by
4 or 5 connate sepaloid bracts. In the axil of each bract develops a group of stamens in a scorpioid
manner. Each stamen represents a naked male flower because it is a jointed structure which indicates
that its upper portion is the filament bearing the anther and its lower portion represents the pedicel
of the male flower bearing stamen. On the rim of the cup-like involucre are present nectar-secreting
glands. Glands are oval or crescent-shaped and often brightly coloured. A cyathium appears like a
single flower but actually it is an inflorescence because (i) naked male flowers (stamens) start rip-
ening centrifugally i.e. from the centre towards periphery, (ii) they are jointed, of which the lower
portion represents pedicel which bears the male flower (stamen), (iii) development of male flower
(stamen) is in a scorpioid manner, and (iv) in the centre is present a single naked female flower.
carpellate oil gland on

flower male flower connate bract
pedicel
bract
nectar gland
male flower
(stamen)
A C
B fruit of 3 fused
carpels
Fig. 21.109 A: Cyathia of Euphorbia helioscopia; B: A cyathium of E. corollata, and C: L.S. cyathium
of E. helioscopia.
Flower Bracteate, usually bracteolate; generally unisexual, monoecious (Baliospermum) or dioecious

(Bridelia), actinomorphic, hypogynous, rarely perigynous (Bridelia).
Floral structures are highly variable, and some aspects are undermentioned:
Perianth Consisting of both calyx and corolla, or only of corolla, or sometimes both are absent;
valvate or imbricate; calyx, when present, consists of 5 parts, usually distinct or united; corolla of
0 to 5 petals, free or united in various ways as under:
(i) In Chrozophora sepals are 5, united and valvate, and petals are 5, free and twisted; (ii) in
Croton, the flowers are often apetalous, and the tepals are distinct in male flowers and usually absent
in female flowers; (iii) in Euphorbia the perianth parts of individual flowers are absent (Fig. 21.110);
(iv) in Jatropha same as in Chrozophora; (v) in Mercurialis sepals are only 3 and united while pet-
als, if present, are free, (vi) in Phyllanthus niruri 6 tepals are present in 2 whorls of 3 each, (vii)
in Ricinus communis male flowers have 5 fused, valvate tepals while the female flowers have 3 to
5 fused, valvate tepals (Fig. 21.111).
Androecium Stamens 1 to 100 or more but usually as many or twice as many as tepals, free or
monadelphous; anthers bilocular, rarely 3 to 4 locular; longitudinal or transversely dehiscent. Stamens
are highly variable in number, position and adhesion, varying from (i) only 1 stamen in each male
flower in Euphorbia, consisting of a pedicel, joint, filament and anther lobes (Fig. 21.110); (ii) 3
stamens in Breynia; (iii) a whorl of 3 monadelphous stamens in Phyllanthus niruri; (iv) 5 stamens
with their profusely branched, tree-like filaments having dithecous, introrse anther lobes at their
apex in Ricinus communis (Fig. 21.111); (v) 8 stamens in Acalypha, (vi) 10 stamens in 2 whorls of
5 each in Jatropha, (vii) 15 stamens in 3 whorls of 5 each in Chrozophora, (viii) 15 to 30 stamens
in Codiaeum, and (ix) 80 to 100 or more stamens in Croton. Stamens are either free or more or less
monadelphous; forming a staminal column by the fusion of the filaments as in Phyllanthus niruri;
in male flowers the disc is usually present in the form of intrastaminal or extrastaminal glands;
sometimes staminodes are present in pistillate flowers.
Gynoecium Usually tricarpellary, rarely bicarpellary (Bridelia, Mercurialis) or pentacarpellary
(Wielandia); syncarpous; superior, rarely semi-inferior (Bridelia); trilocular, one (Ricinus, Euphorbia)
448 Plant Taxonomy
male flower
male
stalk female flower
gland flower gland
ovary
involucre
joint
style
female
flower involucre
stigma
stigma
ovary style L.S. Cyathium
anther A Cyathium joint

lobe
stigma
filament
joint Female Flower
pedicel
pedicel bract
bract
cyathia
Male Flower
leaf
stem
Flowering Branch
Floral Diagram (Cyathium)
Fig. 21.110 Euphorbia pulcherrima Willd. syn. Poinsettia pulcherrima R. Grah.
or two (Phyllanthus) anatropous ovules in each locule; axile placentation; styles 3, each bifurcat-
ing apically into two feathery stigmas; a nectariferous disc is present at the base of the ovary; in
staminate flowers, the gynoecium is sometimes present as a pistillode.
Fruit and Seeds Fruit usually a capsule, but sometimes a drupe. Seeds endospermic, oily, with a
knob-like caruncle (Ricinus communis); embryo straight.
Pollination Cross pollination is necessary because of unisexual flowers. Certain characters (coloured
bracts, well-developed nectaries and glands) also favour insect pollination in some plants.
(a) Male Flower: Br, ≈, , P3+3 or (3–5) or 0, A1–μ or (3–μ), G0 or pistillode.
(b) Female Flower: Br, ≈, , P3+3 or (3–5) or (5), A0 or staminodes, G (3).
branched
stamens stigma
perianth style
ovule
An Open Male Flower ovary

tepal
L.S. Female Flower
Floral Diagram (male flower)

female
flower
unopened
male flower
leaf
anthers
stem
Floral Diagram (female flower) Flowering Branch A Staminal Branch
Fig. 21.111 Ricinus communis L.

Several plants of high economic importance belong to Euphorbiaceae, such as Hevea brasiliensis
(the main commercial source of rubber), Ricinus communis (the source of castor oil), Manihot
esculenta (the source of cassava or tapioca), Emblica officinalis (the source of Amla), Euphorbia
pulcherrima (the famous ornamental Poinsettia), and Codiaeum variegatum (the popular ornamen-
tal plant with variegated leaves, often incorrectly called “Croton”). However, several Euphorbiaceae
are poisonous, causing sickness or death if ingested, or dermatitis if juice contacts the skin. Even
the rainwater dripping from certain plants is enough to cause dermatitis, and therefore should be
handled with utmost care. Some of their specific uses are mentioned below:
• Ornamental Plants: These include Acalypha ciliata, A. hispida, A. wilkesiana, Codiaeum
variegatum (plants with variegated leaves), Croton tiglium (crotons), Euphorbia antiquo-
rum (Tidhara Sehund), E. neriifolia (Sehund), E. pulcherrima syn. Poinsettia pulcherrima
450 Plant Taxonomy
(Poinsettia, with red, pink or white floral leaves), E. splendens (Crown-of-thorns, with
bright red bracts), E. tirucalli (Milk-bush), Jatropha hastata, J. gossypifolia, J. podagrica,
J. panduraefolia, and Trewia nudiflora (False white teak).
• Oil Plants: (i) Croton oil, obtained from the seeds of Croton tiglium (Jamalghota) is used as
a powerful purgative, (ii) Castor oil, obtained from the seeds of Ricinus communis (Arandi)
is mainly used as a vegetable oil and also as a mild laxative, as a lubricant and in paint,
varnish and plastic industries, (iii) Jatropha oil, obtained from the seeds of Jatropha curcas,
is used as a purgative and also in skin diseases, in rheumatism, and also in the manufacture
of soaps, lubricants, candles, etc., (iv) Tung oil, obtained from the seeds of Aleurites fordii,
A. moluccana and A. montana, is used in preparing paints, varnishes, linoleum, India ink
and also in waterproofing the paper, wood, etc.
• Rubber: (i) Over 98% of the total natural rubber, produced in the world, is obtained from
the coagulated latex of Hevea brasiliensis, (ii) Manihot glaziovii (Manicoba rubber) is the
another rubber-yielding plant of this family.
• Cassava or Manioc: Tuberous roots of Manihot esculenta (Cassava) are rich in starch
(Arrowroot) and used for preparing bread, biscuits and other foodstuffs. Bitter cassava con-
tains hydrocyanic acid (HCN) and must be used with due care.
• Medicinal Plants: (i) Emblica officinalis syn. Phyllanthus emblica (Amla) fruits are rich
source of vitamin C, used in treating scurvy, in preparing shampoos, and also in making
the hair dyes (bark and leaves of this tree are used for tanning); (ii) Castor (Ricinus com-
munis) oil is a valued purgative known; (iii) Jatropha gossypifolia leaves are used in eczema,
and roots in leprosy and snake bites; (iv) Entire plant of Synadenium grantii is used as a
stimulant of central nervous system; (v) Croton cascarilla and C. elateria bark is used as a
tonic, (vi) The drug, euphorbium, obtained from the latex of Euphorbia resinifera, is used
as a purgative, (vii) Mallotus philippinensis (Kamela tree) fruits are used as anthelmintic.
• Dyes: (i) Kamela dye, obtained from the fruits of Mallotus philippinensis, is used for
dyeing wool and silk, (ii) Blue dye, obtained from the bark of Jatropha curcas, is used for
dyeing fishing nets, (iii) Purple dye, obtained from the bark of Chrozophora tinctoria is
used in textile industry, (iv) Red dye is obtained from the roots of Kirganelia reticulata.
• Timber Plants: Valuable timber, used for packing cases, tea boxes, veneers, plywood, match
industry and several other similar purposes, is obtained from Aporosa dioica, Bischofia
javanica, Drypetes roxburghii, Gelonium multiflorum, Hemicyclia andamanica, H. elata,
Hura crepitans (sandbox tree) and Trewia nudiflora (False white teak).

Treated under Unisexuales of Monochlamydeae by Bentham and Hooker, Euphorbiaceae was placed
under Geraniales by Engler, and under Euphorbiales by Hutchinson (1959), Takhtajan (1969),
Cronquist (1981) and Thorne (1983). Hickey and King (1988) divided Euphorbiaceae into 4 sub-
families as under:
1. Phyllanthoideae (Cotyledons broader than radicle; ovules 2 per locule; no latex).
2. Euphorbioideae (Cotyledons broader than radicle; ovules 1 per locule; latex present).
3. Porantheroideae (Cotyledons as wide as radicle; ovules 2 per locule; no latex).

4. Ricinocarpoideae (Cotyledons as wide as radicle; ovules 1 per locule; latex present).
Euphorbiaceae resembles Geraniaceae and allied families in the characters of fruit. Characters
such as monadelphous or polyadelphous stamens and carunculate seeds bring it close to Sterculiaceae.
Cronquist opined that Euphorbiales are closer to Sapindales as well as to Geraniales, Linales and
Polygalales, and all these have been derived from Rosaceous-stock (Cronquist, 1981).
21.95 URTICACEAE NETTLE FAMILY


Usually herbs with stinging hairs and watery sap; inflorescence cymose; gynoecium monocarpellary
with single ovule; style one.

Laportea, Boehmeria, Pouzolzia, Sarchochlamys, Urtica, Girardinia, Elatostema.

A family of about 45 genera and 1060 species (Hickey and King, 1988), Urticaceae are distributed
in both tropical and temperate regions. About 12 genera and over 100 species have been reported
from India, mainly distributing in Assam, West Bengal, Kashmir and sub-Himalayan regions. Some
of the largely represented genera along with the number of their approximately reported species are
Pilea (400), Elatostema (200), Boehmeria (100), Cecropia (100), Urtica (50) and Laportea (23).
Urtica (stinging nittle) has sharp, stinging hairs which penetrate in the skin of the animals coming
in their contact and cause sharp burning pain for hours.

General Habit Mostly fibrous herbs (Urtica, Fig. 21.112), infrequently shrubs (Maoutia, Boehmeria),
or rarely small trees (Laportea crenulata); without latex but with watery sap; a few are climbers;
stinging hairs, present in several genera (Urtica, Urera), are long hollow cells with silicified tips
which penetrate the skin like a fine needle; stinging hairs usually absent in several genera (Pilea,
Elatostema, Boehmeria, Maoutia, Parietaria, Cecropia); epidermal cells usually contain cystoliths.
Leaf Simple, alternate or opposite; stipulate but rarely exstipulate (Parietaria); often with stinging
hairs.
Inflorescence Usually cymose with condensed flowers; sometimes catkin-like or heads; or flowers
solitary.
Flower Bracteate or ebracteate, ebracteolate; usually incomplete, unisexual, actinomorphic; rarely
bisexual (Parietaria); hypogynous; usually very small or greatly reduced.
452 Plant Taxonomy
stamen
tepal
A Male Flower Stamen

(before dehiscence) Stamen
stigma (after dehiscence)
ovary
tepal
A Female Flower
Floral Diagram
(male flower)
Flowering Branch
T.S. Ovary
Floral Diagram
(female flower)
Fig. 21.112 Urtica dioica L.
Perianth 4 or 5, free or united tepals; sometimes even tepals are absent (Forskohlea); mostly green
or sepaloid; petals absent.
Androecium In staminate flowers the stamens are mostly 4, rarely 3–5; present opposite the perianth
lobes; straight (Cercopia, Poikilospermum) or bent down inwards in bud and exploding when ripe;
anthers dithecous, dehiscing longitudinally; scale-like staminodes are present in pistillate flowers;
only 1 stamen is present in each male flower in Forskohlea.
Gynoecium In pistillate flowers the gynoecium is monocarpellary; superior; unilocular with 1 basal,
erect, orthotropous ovule; style 1; stigma 1 and often with a brushlike tuft; a rudimentary pistillode
is present in staminate flowers.
Fruit and Seeds Fruit usually an achene or fleshy drupe, enclosed by persistent perianth lobes. Seeds
endospermic, with small and straight embryo.
Pollination Urtica dioica flowers are wind pollinated. The anemophilous flowers of Parietaria dif-
fusa are well-adapted to ensure cross pollination.
(a) Male Flowers: Br or Ebrl, ≈, , P4–5 or (4–5), A4, G0 or pistillode.
(b) Female Flowers: Br or Ebrl, ≈, , P4–5 or (4–5), A0 or staminode, G1.

The family is of little economic importance as under:
• Fibre (ramie), obtained from the inner bark of Boehmeria nivea, is perhaps the longest and
most silky of all vegetable fibres. It is used for cordage, ropes, etc. Fibres obtained from
Maoutia puya and Gigardinia zeylanica are also used for similar purposes.
• Bank notes are manufactured from the paper pulp made from the tough and silky fibres of
Boehmeria nivea.
• Ornamental plants belong to several species of Pilea, Pellionia, Soleirolia, and Urera.
• Young tops of Urtica are cooked and eaten like spinach.
• Fruits of Debregeasia hypoleuca are edible.

Family Urticaceae of Unisexuales (Bentham and Hooker) was placed under order Urticales by most
of the latter taxonomists (Engler and prantl, Hutchinson, Cronquist, Takhtajan, and Thorne). Hickey
and King (1988) divided Urticaceae into 6 tribes (Urticeae, Procrideae, Boehmerieae, Parietarieae,
Forskohleeae and Conocephaleae).
Urticaceae is closely related to Moraceae, but differs from the latter in having erect ovule and a
straight embryo. From the other families of Urticales, Urticaceae are considered advanced for having
herbaceous plants and monocarpellary ovary.
21.96 MORACEAE MULBERRY FAMILY

Monochlamydeae or Apetalae, Unisexuales.1

Shrubs or trees and only rarely herbs; milky sap present; leaves stipulate; flowers unisexual; tepals
usually 4; stigmas 2.
1
Bentham and Hooker did not recognise Moraceae as an independent family but discussed its members under two tribes
(Moreae and Artocarpeae) belonging to family Urticaceae of Unisexuales. However, most of the other workers recognised
Moraceae as an independent family of order Urticales.
454 Plant Taxonomy

Artocarpus, Broussonetia, Ficus, Morus.

A family of 53 genera and over 1400 species (Hickey and King, 1988), Moraceae are distributed
mainly in tropical and subtropical regions of the world, and a few are also found in temperate
regions. Only 15 genera and about 150 species have been reported from India. Major genera along
with the approximate number of their worldover reported species and common names of some of
them are Ficus (800, fig or rubber plant), Dorstenia (170), Artocarpus (47, jackfruit), Maclura (12,
hedge apple), Morus (10, mulberry), and Broussonetia (8, paper mulberry).

General Habit Mostly trees (Ficus benghalensis, F. religiosa, Morus alba), or shrubs with milky
juice, rarely herbs (Dorstenia) or climbers (Ficus benjamina); aerial prop roots grow vertically down-
ward, enter the soil and provide support to the plant in several species of Ficus (F. benghalensis).
Stem Woody, contains milky, sticky latex in long sacs, specially in secondary cortex and phloem
(Ficus elastica, rubber plant).
Leaf Simple, palmately or pinnately veined; alternate; usually with 2 cuducous stipules; petiolate;
leathery; margin entire (Ficus), serrate (Morus) or lobed; on falling, the caducous stipules leave a
semicircular scar in Ficus.
Inflorescence Basic plan is cymose, but the flowers are grouped in variously shaped clusters of
racemes, spikes, umbels, heads, or catkins, or even on the inside of a hollow receptacle i.e. syco-
nium or hypanthodium (Fig. 21.113). Plants may be monoecious or dioecious. In Morus, female
flowers are arranged in pseudospikes while male flowers
in catkins; in Broussonetia the female flowers are arranged apical opening
in pseudoheads while male flowers in pseudoracemes; in
Artocarpus female flowers in pseudoheads while male flow- region with
male flowers
ers in pseudocatkins; in the syconium of Ficus, the male
flowers are arranged towards the ostiole in the hollow cavity
while rest of the cavity is occupied by female flowers, as
in Ficus carica (Fig. 21.113).
Flower Unisexual, plants monoecious or dioecious, incom- region with
female flowers
plete, actinomorphic; pistillate flowers hypogynous; tetram-
erous, small and inconspicuous.
Perianth Usually 4 tepals, free and in 2 whorls of 2 each fleshy receptacle
(Morus), or more or less united, persistent, imbricate or
valvate; tepals usually ovate and hairy; sepaloid; petals
absent.
Androecium Usually 4 stamens, present only in male or
Fig. 21.113 L.S. syconium
staminate flowers, opposite the tepals (Morus), polyandrous,
of Ficus carica.
dithecous, basifixed or versatile; introrse, dehiscence longitudinal; anthers not exploding; only 1–2
stamens in Ficus, Artocarpus; sometimes rudimentary stamens present in female flowers.
Gynoecium Present only in female or pistillate flowers; bicarpellary, syncarpous, ovary superior
(Morus) to inferior (Dorstenia); out of the 2 carpels usually only 1 develops; unilocular, with 1
usually pendulous ovule; style short and bifurcating into 2 coiled stigmas, sometimes rudimentary
pistil is present in male flowers.
Fruit and Seeds Fruit an achene or drupe-like, but usually a multiple fruit developing from the union
of fruits of several different flowers; several achenes develop inside the fleshy receptacle in Ficus;
in Morus several fruits are enclosed in individual fleshy perianth and collectively called a sorosis.
Seeds are with or without endosperm, and with usually curved embryo.
Pollination In Morus, the anthers of the male inflorescence dehisce, and their pollens are carried
by the wind to the prominent paired stigmas of the female flowers. Pollination in Ficus carica is
effected by a female insect (Blastophaga psenes).
(a) Male Flower: Ebr, ≈, , P2+2, A4, G0.
(b) Female Flower: Ebr, ≈, , P2+2, A0, G (2).

Moraceae are important from the economic point of view in providing several fruit plants, orna-
mentals, rubber plant and medicinal plants, as under:
• Fruit plants belonging to Moraceae include Morus nigra (black mulberry or Shahtoot), M.
alba (white mulberry or Tunt, Fig. 21.114), M. serrata (Himalayan mulberry), Ficus carica
(fig or Anjeer), F. palmata (Anjiri), Artocarpus heterophyllus syn. A. integrifolia (jack fruit
or Kathal), A. incisia (bread fruit) and A. lakoocha (monkey fruit or Barhal).
• Certain sacred trees used in religious ceremonies and also for shade include Ficus bengha-
lensis (banyan tree, or Bargad), F. religiosa (Peepal), F. glomerata (Goolar), and F. virens
syn. F. infectoria (Pilkhan).
• Famous ornamental plants include Ficus elastica (Indian rubber plant), Ficus krishnae
(Krishan’s butter cup), Morus nigra, M. alba, Maclura pomifera (Osage orange), and
Dorstenia cordifolia.
• Paper is obtained from the fiber present in the inner bark of Broussonetia papyrifera (paper
mulberry).
• Rubber is obtained from the latex of Ficus elastica (Indian rubber plant) and Castilla elastica
(Panama rubber).
• Leaves of Morus alba and M. nigra are used to feed silkworms.
• Useful timber, utilized in the manufacture of sport goods and several other articles, is
obtained from Morus alba, M. serrata, Ficus sycomorus, Artocarpus altilis, A. chaplasha,
and A. hirsuta.
456 Plant Taxonomy
anther stigma
style
pistillode ovary
tepal
tepal
Male Flower (opened) Female Flower

Floral Diagram (male)
stamen
pistillode
stem
tepal
L.S. Male Flower leaf
fruit
A Female Branch
Floral Diagram (female)
Fig. 21.114 Morus alba L.
• Some plants of medicinal value include (i) Morus alba and M. nigra—bark is used as purga-
tive and vermifuge, (ii) Ficus benghalensis and F. rumphii—latex is used as an anthelmintic,
(iii) Streblus asper (Kurchna)–latex is used as a sedative in neuralgia, and (iv) Antiaris
toxicaria (Upas tree)–latex is used as a cardiac stimulant.
• Fibre obtained from the bark of Ficus articulata and F. semicordata is used for making
ropes.

Moraceae is not recognized as an independent family by Bentham and Hooker. They treated its mem-
bers in tribes Moreae and Artocarpeae under family Urticaceae of Unisexuales. However, almost all
other workers treated Moraceae as an independent family under order Urticales (Engler and Prantl,
1931; Hutchinson, 1959; Takhtajan, 1969; and Cronquist, 1981). Several workers regard Moraceae
as an advanced family of dicotyledons.
21.97 CANNABINACEAE HEMP FAMILY

Monochlamydeae or Apetalae, Unisexuales1.

Aromatic herbs with watery juice; Cannabis is erect while Humulus is climbing; leaves opposite, sim-
ple in Humulus while palmately compound in Cannabis; flowers unisexual, mostly dioecious while
few are monoecious (Humulus japonica); calyx persistent; gynoecium unilocular; fruit achene.
21.97.3 Common Indian Genera

Cannabis, Humulus.

A very small family of only 2 genera (Cannabis and Humulus) and 5 species, Cannabinaceae are
mainly distributed in north temperate regions. Sometimes placed under Moraceae, the Cannabinaceae
differ from Moraceae in possessing 5 floral parts instead of 4. Both the genera of hemp family grow
in India. Cannabis sativa (Fig. 21.115), the only species of this genus, grows wild all over India.
Of the 4 species of Humulus, H. lupulus grows wild in Himalayas and other parts of our country,
H. americanus grows in USA, H. japonica in Japan and H. scandens in China as well as Japan.

General Habit Erect (Cannabis sativa) or climbing (Humulus lupulus) aromatic herbs with watery
juice; female plants of Cannabis sativa are usually larger than the male plants; latex absent.
Leaves Alternate or often opposite leaves, which are simple or palmately compound and pal-
mately veined; stipulate, stipules persistent; petiolate; margins coarsely toothed; venation reticulate
multicostate.
1
Sometimes included in Urticaceae, Lawrence discussed all members of Cannabinaceae under subfamily Cannaboideae
of family Moraceae while Rendle as well as Hutchinson treated all members of Cannaboideae under a separate family
Cannabinaceae. Bentham and Hooker treated all genera of Urticaceae, Moraceae and Cannabinaceae under a single family
Urticaceae of order Urticales. Recently, Cronquist (1981) and Jones and Luchsinger (1988) discussed Cannabinaceae (or
Cannabaceae) under order Urticales of subclass Hamamelidae of class Magnoliopsida (Dicots).
458 Plant Taxonomy
male
inflorescence
stem
leaf
D
A
A Male Branch
A Female Branch stigma

styles
perianth
lobes
styles
C perianth
stamens
E ovary
B
A Male Flower
A Female Flower A Female Flower
(without perianth)
Floral Diagram Floral Diagram

(Female) (Male)
Fig. 21.115 Cannabis sativa L.

Inflorescence In Cannabis, the male plants bear flowers arranged in axillary paniculate cymes
whereas female plants bear flowers arranged in cone-like axillary spikes; in Humulus, the male plants
have mixed panicles of cymes and racemes whereas female plants bear biparous cone-like flowers.
Flowers Flowers unisexual and plants are usually dioecious; usually apetalous and axillary; actino-
morphic; small and inconspicuous; bracteate in Cannabis sativa and bracts are covered by glandular
hairs on their outer surface; the glands on these hairs remain filled with a secretion containing
tetra-hydrocannabinol.
Male Flower
Perianth Staminate flowers arranged in panicles; each flower with a usually 5-parted perianth; lobes
valvate or imbricate, free (Cannabis) or connate at base (Humulus); green.
Androecium Stamens 5, free, opposite the perinath lobes, dithecous, dorsifixed.
Gynoecium Absent.
Female Flower
Perianth Usually absent; if present, then only one perianth lobe enveloping the ovary is present;
colourless.
Androecium Absent.
Gynoecium Monocarpellary, superior, unilocular, one-ovuled ovary; ovule pendulous and curved;
placentation basal; style one with two long arms; stigmas two, minute and simple.
Fruit An achene; usually glandular and surrounded by persistent perianth lobe.
Seeds Endospermic, flattened with curved embryo.
(a) Male Flower: ≈, , P5 or (5), A5, G0.
(b) Female Flower: ≈, , P1, A0, G1.

(a) Cannabis Cannabis sativa (hemp) plants provide a valuable fiber of great strength and
durability. The fiber is widely used for making ropes, twine, carpets, sacks, bags, etc. The finer
grades of this fiber are woven into a cloth that looks like coarse linen. The famous “Jeans” are
prepared from this fiber. The “oil of hemp”, obtained from the seeds of C. sativa, is used for
making varnishes and soft soap. Narcotics (e.g. bhang, ganja and charas) are obtained from the
flowering tops and leaves of C. sativa. The drug hashish or charas, a resinuous substance contain-
ing several powerful alkaloids, is obtained from this plant. Bhang or marijuana is actually the
resinuous substance obtained from the upper leaves, bracts, flowers of the female inflorescences
of C. sativa.
(b) Humulus The inflorescences of Humulus lupulus (Hops) contain a bitter aromatic narcotic
substance called lupulin. Hops are highly used in medicine for their sedative and soporofic
460 Plant Taxonomy
properties. Their principal use, however, is in brewing industry. They are added to beer to prevent
bacterial action and consequent decomposition, and also to improve flavour and to impart the
characteristic bitter taste to the beverage.

As already mentioned in the footnote of Article 21.97.1, the family Cannabinaceae has been treated
variously by different taxonomists i.e. either under Moraceae, or under Urticaceae, or also as an
independent family under order Urticales. But due to its characters like (i) herbaceous or vines with
watery sap, (ii) aromatic plants without stinging hairs, (iii) five floral parts instead of four, (iv) sta-
mens short and straight, (v) ovule apical and anatropous, and (vi) achene type of fruit, Cannabinaceae
should be treated as an independent family.
21.98 CASUARINACEAE CASUARINA FAMILY


Equisetum-like jointed stem; minute, whorled, scaly leaves; unisexual flowers; fruit woody, cone-
like.

It is a monotypic or unigeneric family represented by only one genus (Casuarina) with about 65
species distributed mainly in north-east Australia and also in Fiji, New Caledonia, Malaysia and
Mascarene Islands. Only one species (Casuarina equisetifolia; Fig. 21.116) occurs in India, and this
is the most common species of the world.

Habit Xerophytic, evergreen, shrubs or trees.
Stem Woody, much branched, with jointed, whorled or striate branches having prominent nodes
and internodes; internodes with longitudinal ridges and grooves.
Anatomically, the stem contains (i) ridges and grooves, (ii) thick cuticle, (iii) sunken stomata
in grooves protected by long hairs, (iv) sclerenchyma and green palisade parenchyma in the
ridges, (v) vascular bundles in two alternate rings, (vi) wood contains vessels, tracheids, fibres and
parenchyma.
Leaf Scale-like, 4–16 on each node; whorled; minute, basally connate, forming a sheath around
the node; usually linear to lanceolate; leaf tips appear only as minute teeth; leaves resemble with
that of Ephedra and Equisetum.
male inflorescences
stem
leaves
Pistillate Inflorescence
stigma
Staminate
A Part of Inflorescence
Twig
male style
flowers
ovary
Pistillate
Flower
fruits
Staminate Fruit
Flower
sheath of bracts bracteoles
Flowering perianth lobe stamen

Branch
axis
Floral Diagram
(male inflorescence)
Floral Diagram
(female flower)
Fig. 21.116 Casuarina equisetifolia Forst.

462 Plant Taxonomy
Inflorescence Male flowers arranged in catkin-like erect spikes while the female flowers usually in
spherical heads (Fig. 21.116); in both male and female branches the internodes are short; plants are
monoecious or dioecious.
Flower Bracteate, bracteolate, bracteoles usually 2; incomplete, unisexual; perianth generally absent
or rudimentary.
Male Flower Several male flowers develop at each node of the inflorescence axis; these flowers at
each node remain protected by a sheath formed by the combination of bracts; each male flower is
surrounded by 2 bracteoles, two small tepals or perianth lobes, and a central stamen which usually
hangs out over the edge of the sheath of bracts; anthers dithecous, basifixed, dehiscing by vertical
slits.
Female Flower Several female flowers remain crowded in spherical heads at the ends of the branches;
each female flower is surrounded by a bract, a pair of bracteoles and a centrally located gynoecium;
perianth lobes are absent; gynoecium is bicarpellary, syncarpous; ovary superior, originally bilocular,
but becomes unilocular by the suppression of posterior locule; ovules 2, of which 1 usually aborts;
parietal placentation; style 1, very short; stigmas 2, long, linear and feathery.
Fruit and Seeds Fruit usually a 1-seeded, small-winged samara, surrounded by 2 woody bracteoles
and a bract; several such fruits remain arranged together to form cone-like dry multiple fruit. Seeds
are non-endospermic and the embryo is straight.
Floral Formulae
(a) Male Flower: Br, Brl, ≈, , P2, A1, G0.
(b) Female Flower: Br, Brl, ≈, , P0, A0, G (2).

Casuarina is important mainly as a timber tree. The wood of C. equisetifolia, C. cunninghamia
and C. stricta is used for furniture, for fuel and also for basket making. The bark of C. suberosa
and C. stricta is utilised for tanning. C. equisetifolia is usually grown as an ornamental and also
in order to reclaim sandy dunes in India and several countries.

Treated under Unisexuales by Bentham and Hooker, Casuarinaceae was included under Casuarinales
by Hutchinson (1959), Takhtajan (1969), Cronquist (1981) and Thorne (1983). Several workers believe
Casuarinaceae to be derived from the Hamamelidales (Takhtajan, Thorne, etc.). In the systems of
Engler and Wettstein, Casuarinaceae is placed at the beginning of the dicotyledons. According to
Kuznetsov (1936), Casuarina is one of the simpler types of angiosperm and is “unconnected with
any other family”, and the Casuarinales consist of a single family Casuarinaceae with a single genus
(Casuarina).
On the other hand, Takhtajan (1969) mentioned that because of its special types of leaves and
reproductive structures, Casuarina looks “more like Ephedra than an angiospermous plant”. Takhtajan
listed several characters of Casuarina (such as unisexual anemophilous flowers, unistaminate male
flowers, absence of perianth, reduction of posterior locule in the ovary, etc.) which all “are signs of
high evolutionary development” of this taxon.
In several morphological features Casuarinaceae approaches families such as Urticaceae,
Betulaceae, Corylaceae, Myricaceae and also Hamamelidaceae. “In many features of floral con-
struction, Casuarina undoubtedly comes near to the Urticales” (Takhtajan, 1969). However, the pol-
len grains of Casuarina are very similar to those of Betulaceae and Myricaceae (Erdtman, 1952).
Takhtajan (1969) finally stated that Casuarina comes more closer to Hamamelidaceae because of
similarity in several of their characters, such as ‘unisexual anemophilous flowers, structure of the
infloresence, reduction of perianth, gynoecia of two median carpels and reduction in the number of
ovules”.
21.99 SALICACEAE WILLOW FAMILY

Dicotyledons, Monochlamydeae, Ordines Anomali
Deciduous dioecious trees, shrubs or sub-shrubs; flowers arranged in catkins; each flower subtended
by a bract or scale with a cup-like disc or a gland; seeds with tufts of hairs.

Populus alba (Poplar), Salix tetrasperma (Willow).

Family is represented by only 3 genera (Populus, Salix and Chosenia) and about 530 species, distrib-
uted widely in north-temperate regions of the world. Salix, with about 500 species and Populus with
about 35 species are distributed mainly in north-temperate regions while the single known species
of Chosenia (C. arbutifolia) is found in temperate and sub-arctic regions of north-east Asia.
(i) Salix: Bud remains covered by a single scale or bract; nectaries 1–4; stamens generally 2;
leaves usually short-stalked with narrow blades.
(ii) Populus: Bud remains covered by several imbricate scales or bracts; nectaries absent; stamens
generally 4 to many; leaves usually long-stalked with broad blades.
(iii) Chosenia: Bud in this monotypic genus remains covered by a single scale or bract; nectary
absent; stamens 5.

General Habit Woody deciduous trees or shrubs and sub-shrubs; mostly dioecious and only rarely
monoecious (e.g. Salix medemil); many species are prostrate shrubs containing creeping stem some-
times, half-burried in the soil (e.g. Salix divergens, S. furcata and S. flabellaris). S. nigra and S. fra-
gilisi are tall trees reaching upto 100 feet or more. Poplars are lofty trees attaining a great height.
Stem Aerial, erect or sometimes prostrate, solid, cylindrical, glaucous or glabrous.
464 Plant Taxonomy
Leaves Simple, stipulate; usually alternate or rarely subopposite (e.g. Salix purpurea), deciduous;
petiolate, petiole short (Salix) or long (Populus); ovate to obovate (Salix caprea, Fig. 20.117).
Inflorescence Plants dioecious; flowers arranged in dense erect or pendulous catkins; small bracts
present at the base of the catkins (Salix caprea, Fig. 21.117).
Flower Flowers unisexual, naked, each subtended by a fringed or hairy bract or scale; actinomor-
phic; hypogynous, small, cyclic and inconspicuous; bracts entire (Salix) or toothed (Populus); in
Salix caprea, the unisexual flowers are solitary in the axil of bract and arranged spirally round a
central axis and collectively form catkin-type of inflorescence (Fig. 21.117).
Perianth Absent or vestigial and represented by a cupular disc or small nectary.
Staminate Flowers With 1 or 2 nectariferous glands and 2 to 30 free or basally connate stamens;
in Salix sitchensis the stamens are wholly connate; anthers dithecous, dehiscing vertically.
Pistillate Flowers Bicarpellary to tetracarpellary (Salix tetrasperma), syncarpous, superior, unilocu-
lar; more or less flask-shaped, numerous anatropous ovules on 2 to 4 parietal or basal placentae; 1
or 2 nectariferous glands are present in Salix while absent in Populus; style 1 with 2 to 4 stigmas;
staminodes absent.
Fruit 2 to 4–valved capsule.
Seeds With silky hairs developing from the funicle; hairs not scattered over the entire seed; embryo
straight, endosperm little or absent.
(a) Staminate Flower: Br, ≈, , P0, A2–30, G0.
(b) Pistillate Flowers: Br, ≈, , P0, A0, G (2).

Several species of Salicaceae are grown as ornamentals and also used in basket making and for
pulpwood and floor-boards.
• Salix (Willow): Salicylic acid, the root compound for aspirin, is named for Salix. The soft
inner bark of several species of this plant was widely used as a cure for headache in ancient
times. Aspirin, its chemical derivative, is still used in headache throughout the world. Cricket
bats (S. alba) and polo balls are traditionally made from the wood of willow (Salix). Several
species of Salix are source of wood made into charcoal and twigs used in making baskets.
Some forms of Salix are especially popular as decorative trees in damp situations.
• Populus (Poplar): Besides being well-known for its ornamental value, the bark of some spe-
cies of Populus is also of medicinal value. P. nigra (Lombardy Poplar) has been a familiar
avenue tree for over two centuries. Wood of P. tremula is used for making match-boxes and
match-sticks.
I
A Male Catkin
stamens
B
bract
Male Branch (hairy)
A
nectary
Female Branch
G
A Staminate Flower
ovules
hairs
gynoecium
J
bract
ovary A Fruit
nectary (Dehiscing)
D
C
L.S. Ovary
A Pistillate Flower stigmas
parietal
placentum
ovules
style
H
F A Stamen
E
T.S. Ovary Style and Stigmas
Fig. 21.117 Salix caprea L.

466 Plant Taxonomy

According to Lawrence, there is much evidence that Salicaceae “has been derived from advanced
ancestral stocks but data are yet needed before the phyletic position can be indicated”. Hutchinson
(1926) retained Salicaceae with other amentiferous families of presumed hamamelidaceous affinities
and treated it as the most primitive of them. Lawrence mentioned that “there is evidence to indicate
the Salicaceae to be a more highly specialized group than most of the other amentiferous families”
and the only one of an order (Salicales) that stands well apart from other orders of dicotyledons.
Bentham and Hooker included 4 families (Salicaceae, Lacistemaceae, Empetraceae and
Ceratophyllaceae)under the series Ordines Anomali of Monochlamydeae and regarded all of them as
advanced families. Benson (1957), however, included Salicaceae under order Salicales of Amentiferae
and regarded it as an advanced family. Workers like Hutchinson (1973), Takhtajan (1980) and
Cronquist (1981) treated Salicaceae under order Salicales while Thorne (1983) treated it under order
Violales.
Salicaceae is related to Juglandaceae, Myricaceae and Fagaceae in possessing unisexual flowers
arranged in catkins and their perianth is represented simply by glands or scales. It, however, differs
from these families in possessing capsular fruits with numerous hairy seeds and not the drupe or
nut borne by these families.

1. Bentham and Hooker divided Dicotyledons into three subclasses. What are these subclasses?
How can these subclasses be differentiated?
2. Write the characteristic features of the androceium and gynoceium of Magnoliaceae and
Annonaceae.
3. Describe family Ranunculaceae or Papaveraceae in semitechnical language.
4. Write the name of the family of following genera:
1. Iberis 2. Coronopus 3. Cleome 4. Fumaria
5. Argemone 6. Eschscholzia 7. Nelumbo 8. Delphinium
9. Nigella 10. Clematis 11. Annona 12. Michelia
5. Describe the economic importance of family Malvaceae or Tiliaceae.
6. Citrus fruits belong to which family? Describe the Citrus family in semitechnical language.
Also write a note on its economic importance.
7. Name any three families of your choice belonging to Disciflorae. Describe details of any
one of them in semitechnical language.
8. What are the three subfamilies of family Leguminosae. Make a key to differentiate between
these three subfamilies.
9. Describe in detail the economic importance of Papilionoideae.
10. “Rosaceae show great diversity in types of fruit”. Justify this statement. Also write a note
on the edible fruits of this family.
11. Following genera belong to series Calyciflorae of Polypetalae. Name the family they belong
to:
1. Daucus 2. Foeniculum 3. Mammillaria 4. Passiflora
5. Begonia 6. Luffa 7. Momordica 8. Lagenaria
9. Lawsonia 10. Eucalyptus 11. Callistemon 12. Terminalia
13. Pyrus 14. Prunus 15. Dalbergia 16. Bauhinia
12. Give the floral description of Cucurbitaceae or Umbelliferae (Apiaceae).
13. Name all the three series of Gamopetalae proposed by Bentham and Hooker. Describe the
floral structure of any of the families of your course belonging to Gamopetalae.
14. Describe in detail the inflorescence and detailed floral structure of Asteraceae.
15. Compositae (Asteraceae) occupy highest position among angiosperms. Comment on this
statement.
16. Describe family Asclepiadaceae in semitechnical language.
17. Write the botanical name and the family of following plants:
1. Sarpgandha 2. Karonda 3. Chiku 4. Coffee
5. Coriander 6. Ajwain 7. Heeng 8. Zeera or cumin
9. Saunf 10. Kheera
18. Describe family Convolvulaceae or Solanaceae using semitechnical language.
19. Compare the floral structures of Acanthaceae and Lamiaceae.
20. Give characteristic features of all the eight series of Monochlamydeae. Give at least one
example of family of each of these series.
21. Describe the floral structure of family Euphorbiaceae.
Suggested Reading
Bentham, G. and J.D. Hooker, 1862–1883, Genera Plantarum, 3 Vols., Reeve & Co., London.
Britton, N.L. and J.N. Rose, 1919–1923, The Cactaceae, Vols. 1–4, The Carnegie Institution, Washington.
Cronquist, A., 1981, An Integrated System of Classification of Flowering Plants, Columbia Univ. Press,
New York.
Engler, A.W. and K. Prantl, 1887–1915, Die Naturlichen Pflanzenfamilien, 23 Vols., Leipzig.
Heywood, V.H., 1978, Flowering Plants of the World, Mayflower Books, New York.
Hickey, M. and C. King, 1988, 100 Families of Flowering Plants, (2nd ed.), Cambridge University Press,
Cambridge.
Holmes, S., 1983, Outline of Plant Classification, Longman, London.
Hutchinson, J., 1969, Evolution and Phylogeny of Flowering Plants, Academic Press, London.
________ 1973, The Families of Flowering Plants, (3rd ed.), Clarendon Press, Oxford.
Jones, S.B. Jr. and A.E. Luchsinger, 1987, Plant Systematics, (2nd ed.), McGraw-Hill Book Co., New
York.
468 Plant Taxonomy
Lawrence, G.H.M., 1951, Taxonomy of Vascular Plants, Macmillan Co., New York.
Radford, A.E., 1986, Fundamentals of Plant Systematics, Harper & Row Publishers, Inc., New York.
Rendle, A.B., 1925, The Classification of Flowering Plants Vol. II (Dicotyledons), Cambridge Univ. Press,
London.
Takhtajan, A., 1969, Flowering Plants: Origin and Dispersal, Oliver & Boyd Ltd., Edinburgh.
________ 1980, Outline of the classification of flowering plants (Magnoliophyta), Bot. Rev. 46: 225–359.
Thorne, R.F., 1983, Proposed new realignments in the angiosperms, Nordic J. Bot. 3: 85–117.
Tutin, T.G., 1964, Flora Europaea Vol. I, Cambridge Univ. Press, London.
Willis, J.C., 1973, A Dictionary of the Flowering Plants and Ferns, (8th. ed., revised by H.K. Airy Shaw),
Cambridge Univ. Press, London.
C
H
A
P
T
E
R
SELECTED
FAMILIES OF
MONOCOTYLEDONS 22
22.1 MONOCOTYLEDONS AND THEIR CLASSIFICATION
Monocotyledons (usually characterised by the presence of one cotyledon, fibrous and adventitious
roots arising from the base of the stem, narrow leaves with parallel veins, herbaceous stem, irregu-
lar distribution of vascular bundles in the stem, trimerous flowers, and perianth often not clearly
divisible into calyx and corolla) were also divided into varying number of families by different
workers, similar to dicotyledons. Bentham and Hooker (1862–1883) recognised 34 families under
Monocotyledons while Takhtajan (1969) divided class Liliatae (or Monocotyledons) into 69 families.
Cronquist (1981) included 65 families under class Liliopsida (Monocots) whereas Thorne (1983)
treated Monocotyledoneae (= Liliidae) as a subclass of class Angiospermae (= Annonopsida) and
discussed 53 families under this subclass. Only 16 monocot families are discussed in the present
chapter. The criteria of selecting the discussed families, compilation of their characters, method of
family description, and their arrangement in this chapter, are almost the same as followed in Chapter
21 (Selected Families of Dicotyledons).
Bentham and Hooker divided Monocotyledons into 7 series as under:
Series 1. Microspermae, characterised by the presence of epigynous flowers, inferior ovary,
parietal placentation and very small and numerous non-endospermic seeds. It includes 3 families,
namely Hydrocharitaceae, Orchidaceae1 and Burmanniaceae.
Series 2. Epigynae, characterised by epigynous flowers, inferior ovary, and large endosper-
mic seeds. It includes 7 families, namely Iridaceae, Amaryllidaceae1, Haemodoraceae, Taccaceae,
Dioscoreaceae, Bromeliaceae1 and Scitamineae1.
Series 3. Coronarieae, characterised by coloured or petaloid perianth, superior ovary and endosper-
mic seeds. It includes 8 families, namely Liliaceae1, Commelinaceae1, Pontederiaceae, Rapatiaceae,
Xyridaceae, Roxburghiaceae, Philyderaceae and Mayaceae.
Series 4. Calycineae, characterised by green or sepaloid perianth, superior ovary and endospermic
seeds. It includes 3 families, namely Juncaceae1, Palmae1, and Flagellariaceae.
1
Discussed in the present text.
470 Plant Taxonomy
Series 5. Nudiflorae, characterised by perianth mostly lacking or present in the form of scales or
bristles, superior ovary and seeds endospermic. It includes 5 families, namely Typhaceae1, Araceae1,
Pandanaceae, Lemnaceae and Cyclanthaceae.
Series 6. Apocarpeae, characterised by free carpels, superior ovary and non-endospermic seeds.
It includes 3 families, viz. Alismaceae1, Najadaceae, Triuridaceae.
Series 7. Glumaceae, characterised by small, scale-like or chaffy perianth or no perianth, large
scaly bracts, flowers in spikelets or heads, ovary unilocular with one ovule in locule, and seeds
with abundant and starchy endosperm. It includes 5 families namely Cyperaceae1, Eriocaulaceae,
Restionaceae, Centrolepidaceae and Gramineae1.
22.2 ORCHIDACEAE ORCHID FAMILY

Monocotyledons, Microspermae, Orchidaceae.

Perennial herbs; perianth inconspicuous; leaves 2-ranked; flowers zygomorphic, usually trimerous;
usually labellum, pollinia and gynostegium present; ovary inferior.

Cattleya, Cymbidium, Cypripedium, Dendrobium, Habenaria, Orchis, Vanda, Zeuxine.

Orchidaceae is one of the largest families of the flowering plants, represented by about 1000 genera
and 20,000 species which are cosmopolitan but primarily distributed in tropical areas. Its members
are rare in arctic regions. In Indian flora this is the second largest family represented by about 130
genera and over 880 species, distributed mainly in Eastern Himalayas, Western Ghats and Khasi
hills.
This is the family of greatest variety of flowers among flowering plants. Orchid flowers are known
for their strange shape, longevity and beautiful look. Some of the largely represented genera along
with the number of their approximately reported species are Dendrobium (1500), Habenaria (600),
Epipendrum (400), Oncidium (350), Odontoglossum (200), Cattleya (60), Vanda (60), Cypripedium
(50) and Cymbidium (40).

General Habit Perennial herbs; either terrestrial (Orchis), epiphytic (Cattleya, Cypripedium) or sap-
rophytic (Neottia); orchids in temperate regions are mainly terrestrial while those in tropical regions
are mainly epiphytic; sometimes climbers (Bulbophyllum, Vanilla); often with fibrous or tuberous
roots or rhizomatous; epiphytic orchids usually contain fleshy pseudobulbs consisting of one or more
1
Discussed in the present text.
Selected Families of Monocotyledons 471
petal
sepal
stigmatic lobes
column
dorsal sepal
A E
lip
lateral petal
viscidium
pairs of
rostellum
pollinia
column
caudicle
labellum
united lateral
sepals
ovary
G
D
C
lip
anther
style
ovary
F
nectary stigmatic
stylar canal
cavity
Fig. 22.1 A–F. Floral details of Epidendrum tampene–A: A flower; B: Column; C: Column with
anther removed; D: Top of column with anther lifted; E: A pair of pollinia; F: L.S. flower; G: Flower of
Cypripedium.
thickened internodes of stem; aerial roots often contain velamen; mycorrhiza often present; stems
leafy or scapose, may be sympodial (Dendrobium) or monopodial (Vanda); saprophytic forms usu-
ally devoid of chlorophyll (Corallorhiza, Neottia).
472 Plant Taxonomy
Leaf Simple, alternate, often distichous, rarely opposite; sometimes reduced to achlorophyllous
scales (e.g. saprophytic species); usually fleshy and linear to ovate in shape, often encircling the
stem with a sheathing base; foliage leaves absent in some species of Dendrophylax and Epipogon;
only a single leaf per shoot is present in Pleurothallis.
Inflorescence Variable from racemose, paniculate (Oncidium), spicate, or flowers solitary
(Cypripedium).
Flower Usually bracteate; bisexual or rarely unisexual; zygomorphic; epigynous; showy, beautifully
coloured; and extraordinarily diverse in shape, size and colour; mostly resupinate i.e. twisted 180°
or upside down.
Perianth Usually 6 tepals in 2 whorls of 3 each; the outer whorl (representing calyx) of 3 tepals is
green or coloured; the inner whorl (representing corolla) of 3 tepals is petaloid, of these the posterior
tepal is highly modified, often projected basally into a spur and called labellum or lip.
Labellum is strictly the uppermost petal but looks as if located on the lower side of the flower in
most orchids. This is caused either by the twisting of the ovary through 180° (as in many-flowered
orchids), or by the pedicel bending back over the apex of the stem (as in single-flowered orchids).
This phenomenon is called resupination. It enables the labellum to work as a landing-place for pol-
linating insects.
Androecium Represented by 2 fused lateral stamens or 1 terminal stamen; anthers dithecous,
introrse; pollen grains granular or bound together by viscin threads into masses called pollinia.
Stigmas, style, and stamens are adnate to form a single, highly complex structure called column
or gynandrium (Fig. 22.1), which represents the most characteristic part of the orchid flower.
In several orchids (Zeuxine, Fig. 22.2) a pair of pollinia remain connected to the glandula or
corpusculum with the help of a stalk called caudicle. A connection between the ovary and stamen
is made by rostellum. Sometimes staminodes are also present.
Gynoecium Tricarpellary, syncarpous, inferior, unilocular, many ovules in the locule; parietal pla-
centation; rarely the ovary is trilocular and the placentation is axile (Apostasia); style 1 (part of the
column); stigmas 3, of which 2 lateral are often fertile and the third one is sterile forming a beak
called rostellum.
Fruit and Seeds Fruit usually a capsule, containing a very large number of very small, non-endosper-
mic seeds, which are distributed easily by wind.
Pollination Flowers of Orchidaceae are well-adapted for insect pollination because of several char-
acters, such as large-sized and gaudy perianth, pleasant smell, and presence of nectar secreted in
nectary or sac-like rostellum. The flowers are protandrous. When an insect enters the flower, the
pollinia get attached on its body by their caudicles. When this insect visits another flower, it may
deposit the pollen grains on its stigmatic surface below the rostellum, thus completing the process
of pollination.
General Floral Formula Br, , , P3+3, A(2) or 1, G (3).
anther
perianth bract
labellum
bract labellum
rostellum
caudicle
A Flower stigma
column
ovule
ovary wall
ovule ovary
pedicel
ce
L.S. Flower
cen
res
Inflo
T.S. Ovary
glandula
leaf
caudicle
stem
pollinium root
Pollinia
Flowering Plant Floral Diagram
Fig. 22.2 Zeuxine strateumatica L.

• The fruits of the climbing genus Vanilla fragrans syn. V. planifolia are the source of highly
fragrant extract used in perfumery and confectionery.
• Tuberous roots of Habenaria susannae and Orchis latifolia are edible and eaten during
scarcity.
474 Plant Taxonomy
• Starchy tubers of some genera (Cymbidium, Dactylorhiza, Eulophia, Orchis, etc.) are dried
to form “Salep” of commerce, used for culinary and medicinal purposes.
• Orchids are known throughout the world for their beautiful flowers of various forms,
shapes and highly attractive colours. Some orchid genera commonly grown in glasshouses
are Cattleya (florist’s orchid), Cymbidium, Cypripedium (lady’s slipper), Dendrobium,
Epidendrum (green-fly orchid), Habenaria (fringe orchid), Miltonia, Odontoglossum (baby
orchid), Oncidium (butterfly orchid), Orchis (showy orchid), Paphiopedalum, Phalaenopsis,
Vanda, Vanilla and Zeuxine.

Treated as a family under series Microspermae by Bentham and Hooker, Orchidaceae was placed
under a separate order Orchidales by majority of the later workers including Hutchinson (1959),
Takhtajan (1969) and Cronquist (1981). Thorne (1983), however, placed Orchidaceae under the
suborder Orchidineae of the order Liliales. Willis (1973) divides Orchidaceae into 3 subfamilies
(Apostasioideae, Cypripedioideae and Orchidoideae) divisible further into 6 tribes (Apostasieae,
Cypripedieae, Orchideae, Neottieae, Epidendreae and Vandeae).
Several taxonomists consider Orchidaceae to be the most advanced and highest evolved among
monocotyledons. The characters which support this view include (i) reduction in number of stamens,
(ii) resupinate epigynous ovary, (iii) presence of rostellum, (iv) non-endospermic seeds, (v) herbaceous
habit, and (vi) presence of several epiphytes.
According to the majority of taxonomists, Orchidaceae originated from a Liliaceous stock.
Hutchinson opined that Orchidaceae originated from Liliaceae through Hypoxidaceae and
Apostaciaceae. Some, however, also trace another line of parallel evolution of Orchidaceae from
Musaceae.
22.3 IRIDACEAE IRIS FAMILY

Monocotyledons, Epigynae, Iridaceae.

Perennial herbs often with bulbs, corms, or rhizome; contain equitant leaves; flowers trimerous with
3 petaloid sepals and 3 petals; stamens 3; ovary inferior.

Belamcanda, Crocus, Gladiolus and Iris.

A family of about 70 genera and over 1500 species, Iridaceae are distributed in both tropical and
temperate regions with the chief centres of distribution in tropical America and South Africa. Chief
areas of distribution in India are Kashmir and Kumaon hills. Common species cultivated in Indian
gardens include Belamcanda chinensis, Crocus sativus, C. biflorus, C. speciosa, Gladiolus primuli-
nus, G. colvillei, Freesia refracta, Iris kumaonensis, I. germanica and I. foetidissima. Iridaceae are
cultivated throughout the world for their beautiful flowers.

General Habit Perennial herbs with roots produced from corm (Crocus, Fig. 22.3A, B), bulb
(Gladiolus), or rhizome (Iris, Fig. 22.3C; Belamcanda); very rarely subshrubs; stems solitary or
several, or plants scapose.
Leaf Usually simple, exstipulate, sessile, linear to ensiform, equitant in 2 ranks; mostly basal,
numerous and crowded at the base of stem; with sheathing leaf base; parallel venation.
Inflorescence Racemose raceme or paniculate, or flowers solitary (Sisyrinchium, Romulea); invari-
ably terminal; in several genera the flowers develop on distinct aerial flowering axis or scape
(Iris).
Flower Bracteate, complete; actinomorphic or zygomorphic (Gladiolus, Tritonia); bisexual, epigy-
nous, trimerous; showy, large and beautifully coloured; usually subtended individually or in groups
by 2 spathe-like bracts.
Perianth Usually 6 tepals arranged in 2 whorls of 3 each; of these the outer whorl of 3 tepals
represents calyx and often petaloid; and inner whorl of 3 tepals represents corolla; tepals of both
the whorls are often distinguishable from each other by colour, size, or texture; free or united, but
usually basally connate into a long tube (Crocus).
Androecium Usually 3 distinct stamens situated opposite the tepals of the outer whorl; mostly epi-
phyllous; anthers dithecous, basifixed, extrorse, dehiscing by vertical slits.
Gynoecium Tricarpellary, syncarpous; ovary inferior, trilocular, with few to many anatropous ovules
in each locule, axile placentation; in Hermodactylus the ovary is unilocular with parietal placenta-
tion; style 1, often 3-branched, long and coloured; stigmas 3, filiform to subulate or fimbriate; ovary
rarely superior (Isophysis); style is very short in Eleuthrine.
Fruit and Seeds Fruit a loculicidal capsule dehiscing by 3 valves; seeds with small embryo and
copious endosperm, sometimes arillate.
Pollination Usually entomophilous, effected by bees and butterflies.
General Floral Formula Br, ≈ or , , P(3+3) or (3+3), A3 , G (3).

• Dried styles of Crocus sativus are used to make ‘saffron’, an orange-yellow dye used chiefly
for flavouring and colouring dishes. Saffron is also used to give colour to the cooked rice.
“Around 100,000 styles are required to produce 1 kilogram of saffron” (Jones and Luchsinger,
1987).
• Orris root, used in perfumes and in dentifrices, is obtained from the rhizomes of Iris ger-
manica var. florentina.
476 Plant Taxonomy
stigma tepal
flower
stamen
inner perianth segment
style
outer perianth
segment
stigma
bract
stamen
perianth tube
stylar column
B ovary
spathe
scape
scape
leaves D
corm
rhizome
roots
A C
Fig. 22.3 Entire plant (A) and L.S. of the upper part of flower (B) of Crocus tomasinianus; Entire plant
(C) and L.S. flower (D) of Iris pseudacorus.
• Roots of Belamcanda chinensis are an antidote to snake poison. A drug obtained from its
rhizomes is used in tonsilitis.
• Roots of Iris germanica are diuretic and used in diseases of gall bladder.
• Iridaceae are highly-prized decorative plants of gardens and parks. Some of the ornamental
genera are Belamcanada (blackberry lily), Crocosmia, Crocus, Dierama, Freesia, Gladiolus,
Iris, Ixia, Romulea, Sisyrinchium (blue-eyed grass), Tigridia (tiger flower) and Tritonia.

Hutchinson (1959) and Takhtajan (1969) placed Iridaceae under order Iridales while Cronquist
(1981) and Thorne (1983) treated it under order Liliales. Engler and Prantl included Iridaceae under
Liliiflorae and divided it into 3 tribes viz. Sisyrinchieae, Ixieae and Irideae, and the same classifica-
tion has been followed by Hickey and King (1988).
Iridaceae is considered to be close to Liliaceae and Amaryllidaceae. Hutchinson traced its ori-
gin from Liliales while Takhtajan opined that Iridales “probably derived directly from the family
Liliaceae, most likely through subfamily Melanthioideae”.
22.4 AMARYLLIDACEAE DAFFODIL FAMILY

Monocotyledons, Epigynae, Amaryllidaceae.

Bulbous or rhizomatous, perennial herbs; inflorescence usually a leafless scape or umbel subtended by
spathe-like bracts; perianth gamophyllous; stamens 6, antiphyllous and epiphyllous; ovary inferior.

Agave, Crinum, Narcissus, Zephyranthes.

A family of about 85 genera and over 1100 species (Hickey and King, 1988), Amaryllidaceae are
usually distributed in tropical and subtropical regions. About 9 genera and over 60 species have been
reported from India. Some of the larger genera along with the number of their worldover reported
species are Crinum (110), Hippeastrum (75), Narcissus (60), Hymenocallis (50), Haemanthus (50),
Cyrtanthus (47), Zephyranthes (40), Galanthus (20), and Pancratium (15).

General Habit Usually perennial, bulbous (Crinum) or rhizomatous (Hypoxis), xerophytic herbs
leafing only in the rainy or spring season; some are arborescent (Agave, Furcraea).
Leaf Mostly basal, simple, alternate, often distichous, exstipulate, sessile with sheathing leaf base
(Crinum); very large, more or less linear, strap-shaped or ensiform; entire; parallel venation.
Inflorescence Usually on a long leafless scape, with one or more spathes or bracts, cymose, but gen-
erally condensed in the form of an umbel or head; sometimes solitary or paired flowers are present
(Narcissus, Fig. 22.4; Zephyranthes); inflorescence appears after several years in Agave.
Flower Bracteate, complete, bisexual; actinomorphic (Crinum, Fig. 22.5) or zygomorphic (Amaryllis);
trimerous, epigynous; large, showy and usually fragrant.
478 Plant Taxonomy
outer perianth stigma

segment
corona
stamen
inferior ovary
style
perianth
tube
spathe
scape
Fig. 22.4 L.S. flower of Narcissus. Fig. 22.5 Floral diagram of Crinum asiaticum.
Perianth Tepals 6, arranged in 2 whorls of 3 each, not differentiated into calyx and corolla; petaloid;
free or often united basally to form a long tube; salver-shaped or funnel-shaped (Crinum); large and
showy; valvate or twisted or imbricate; sometimes corona present (Narcissus) and looks like an extra
whorl of perianth-segments in between the normal whorl of perianth and stamens.
Androecium Stamens 6, arranged in 2 whorls of 3 each, epiphyllous; present opposite the tepals;
inserted at the throat of perianth tube; filament usually long and coloured (Crinum); anthers dith-
ecous, basifixed or versatile (Crinum); introrse; dehiscence by longitudinal slits or rarely by apical
pore (Galanthus); sometimes some stamens become staminodial.
Gynoecium Tricarpellary, syncarpous; ovary inferior, rarely half-inferior; trilocular or rarely uni-
locular (Alstroemeria); numerous anatropous ovules in each locule; axile placentation; style 1; stigma
capitate or trilobed.
Fruit and Seeds Fruit a loculicidal capsule or berry (Haemanthus). Seeds with small and straight
embryo, and endospermic.
Pollination It is entomophyllous. Insects are attracted because of gaudy-coloured, often scented
perianth and presence of nectar. Nectar is secreted in the inner tepals in Galanthus and in corona
in Narcissus, Hippeastrum, etc.
General Floral Formula Br, ≈ or , , P3+3, A3+3 , G (3).

• Common ornamental plants of Amaryllidaceae are Agave americana (century plant), A. mex-
icana, A. vera-cruz (Kuwarbuti), Alstroemeria aurantiaca, Amaryllis belladona (Amaryllis),
Chlidanthus fragrans, Clivia miniata (Kafir-lily), Crinum asiaticum (Sukhdarsan), C.
defixum, C. latifolium, Curculigo capitulata, C. latifolia, Eucharis grandiflora (Amazon lily),
Galanthus nivalis, Haemanthus coccineus (blood lily), Hippeastrum vittatum, Hymenocallis

americana (spider lily), Hypoxis hirsuta (stargrass), Leucojum vernum (Snowflake),
Narcissus pseudo-narcissus (Daffodil or Nargis), N. bulbocodium, N. tazetta, Nerine filifolia,
Pancratium maritinum, Polianthes tuberosa (Tuberose), Zephyranthes grandiflora (Zephyr
lily), Z. candida and Z. tubispatha.
• The oil obtained from the flowers of several species of Narcissus and Polianthes are used
in high grade perfumery.
• The fibres obtained from Agave cantala, A. sisalana (Sisal hemp), A. vera-cruz and Furcraea
foetida (Mauritius hemp) are used for making cordage, mats, socks, shoe soles, ropes, twines,
etc. The fibres obtained from the leaves of Curculigo latifolia are used for making fishing
nets.
• Fruits of Curculigo latifolia and roots of C. orchioides and Alstroemeria are ground and
eaten as flour.
• Tubers of Curculigo orchioides are used in the treatment of asthma and jaundice.

Treated under series Epigynae by Bentham and Hooker along with other families with inferior ovary,
Amaryllidaceae is included variously by different workers under Liliiflorae (Engler and Prantl),
Amaryllidales (Hutchinson) and Liliales (Takhtajan).
Most authors treat Amaryllidaceae very close to Liliaceae. Hutchinson (1973) opined that the
character of inferior verses superior ovary has been overemphasized in the monocotyledons, although
he treated Amaryllidaceae and Liliaceae as separate families. Cronquist (1981), however, considers
the separation of Amaryllidaceae and Liliaceae on the basis of ovary position to be unreasonable
and merges these two families together under Liliaceae, and the same treatment is followed by Jones
and Luchsinger (1987). Takhtajan (1969) stated that “Amaryllidaceae probably have a common origin
with Agavaceae from Liliaceae”.
22.5 BROMELIACEAE PINEAPPLE FAMILY

Monocotyledons, Epigynae, Bromeliaceae

Short-stemmed, herbaceous, epiphytic plants, often showing xerophytic habit; stem reduced; leaves
stiff, often brightly coloured towards the base; presence of multicellular or stellate scales or hairs;
flowers trimerous and floral bracts usually coloured; calyx herbaceous; anthers versatile.

Ananas, Bromelia, Billbergia, Tillandsia.
480 Plant Taxonomy

Pineapple family is represented by about 45 genera and over 2000 species, and distributed chiefly
in tropical America and West Indies. The chief genera along with their approximately recorded spe-
cies are Tillandsia (500), Pitcairnia (250), Vriesea (190), Aechmea (150), Puya (120), Dyckia (80),
Navia (60), Billbergia (50), Bromelia (40) and Ananas (5). The delicious fruit of “pineapple” comes
from this family. Its bracts, pedicels, ovaries and stalk develop and fuse into a sweet, juicy mass.
A familiar epiphyte of this family is Tillandsia usneodes (Spanish moss).

General Habit Mostly epiphytes (Billbergia nutans, Fig. 22.6), often xerophytic in habit, short-
stemmed.
Leaves Reduced stem contains a basal rosette of linear and spinulose fleshy leaves showing a close
fitting at the base, thus providing the whole plant the shape like that of a funnel; minute scaly hairs
cover the base of the leaves; usually stiff and often spiny; brightly coloured towards the base.
Inflorescence A terminal raceme, panicle, spike or head with flowers developing in the axils of
often brightly-coloured bracts.
Flowers Bracteate; actinomorphic or weakly zygomorphic; usually bisexual, rarely functionally
unisexual; epigynous to hypogynous.
Perianth 6 tepals in 2 series, of which outer 3 calyx-like free parts and inner 3 corolla-like free or
variously connate parts; often brightly coloured; scales or prominent nectaries are present in some
members within perianth lobes.
Androceium Stamens 6, often epipetalous and develop mostly at the base of perianth lobes; free
or partially fused to them; anthers dithecous, linear, usually versatile (Billbergia nutans, Fig. 22.6);
introrse.
Gynoceium Tricarpellary, syncarpous; ovary inferior, half-inferior, or superior; trilocular; numer-
ous anatropous ovules in each locule on axile placentation; style 1; stigmas 3, spirally twisted
(Fig. 22.6).
Fruit A berry, capsule, multiple type (e.g., sorosis in pineapple); often more or less enveloped by
the persistant perianth.
Seeds Very light, sometimes winged; with small embryo and mealy endosperm.
General Floral Formula Br, ≈, , P3+3 or (3+3), A6 , G (3).

• Bromeliaceae is known throughout the world for being the family of famous and delicious
fruit “pineapple” (Ananas comosus). Its leaves are used for making “pina-cloth”.
filaments of
stamens
inflorescence
style
D
L.S. Flower
(Lower Portion)
stigmas
(3)
A
E style stigma C
A stamen
Style (Upper part) (Upper Part)
A Mature Flowering Plant
style
petals
stamen (3)
style
perianth
locules lobes
(6)
ovules sepals
(3)
ovules
ovary
G
B
T.S. Ovary F
L.S. Ovary
A Bisexual Flower
Fig. 22.6 Billbergia nutans H. Wendl.

482 Plant Taxonomy
• Several species of a number of genera of this family are grown as indoor plants for ornamen-
tal purposes. Some of these genera include Billbergia, Cryptanthus, Nidularium, Tillandsia
and Vriesea.
• “Spanish moss” (Tillandsia usneoides) is grown in greenhouses as a curiosity because it
appears like that of a lichen and grows pendent.
• Some species, including T. usneoides, are used as packaging material and also in
upholstery.
• Several species of Bromeliaceae are sources of cordage and fiber for fabrics.

Bromeliaceae has been included under series Epigynae along with six more families, namely
Iridaceae, Amaryllidaceae, Haemodoraceae, Taccaceae, Dioscoreaceae and Scitamineae. Engler
and Prantl included Bromeliaceae under order Farinosae along with 12 more families. Modern
workers like Cronquist (1981) treated Bromeliaceae under a separate order Bromeliales of subclass
Zingiberidae of class Liliopsida while Dahlgren (1983) discussed it under order Bromeliales of
superorder Bromelliflorae of subclass Liliidae (Monocotyledoneae).
Following Engler and Prantl, Hickey and King (1988) classified Bromeliaceae under 4 subfami-
lies, including (i) Navioideae (with spinulose-dentate leaves, superior ovary, capsule fruit and seeds
without wings, e.g. Navia), (ii) Pitcairnioideae (with entire leaves, superior or half-inferior ovary, and
appendaged seeds, e.g. Pitcairnia), (iii) Tillandsioideae (with entire leaves, superior ovary and hairy
seeds, e.g., Tillandsia), and (iv) Bromelioideae (with spinulose leaves, inferior ovary, fruit berry and
naked seeds, e.g. Ananas, Bromelia). From the phylogeny view point, Bromeliaceae represents “the
climax of a line of descent wherein the calyx and corolla have remained distinct or fairly distinct
from each other, a feature retained from the Dicotyledonous stock” (Hutchinson, 1934). Hutchinson
treated Bromeliaceae as related to Commelinales.
22.6 CANNACEAE CANNA FAMILY

Monocotyledons, Epigynae (Treated by Bentham and Hooker under Scitamineae).

Large, coarse, perennial herbs with a tuberous rootstock; leaves large, foliaceous, spirally arranged,
with its petiole sheathing the stem; inflorescence raceme or panicle or 2-flowered cincinni; flowers
large zygomorphic; usually only 1 functional stamen and remaining 4 staminodes; inferior ovary;
ligule absent.

Canna.

Cannaceae is a family of the single genus, Canna (Fig. 22.7), with about 55 species. This famous,
large-flowered ornamental is distributed widely in tropical and subtropical regions of America, Africa
and Asia, including India.
Cannaceae can be distinguished from Marantaceae by the absence of a pulvinus and from the
Zingiberaceae by the absence of a ligule.

Habit Canna is a perennial herb perennating by means of rootstock which is a tuberous rhizome.
Leaves Cauline, large and foliaceous, oblong to broadly elliptical in shape; pinnately veined with
a well-developed midrib; petiolate, petiole sheathing the stem; ligule absent.
Inflorescence Racemose raceme or panicle, made up of 2-flowered cincinni.
Flowers Large, showy, bracteate, bisexual, zygomorphic, shortly-pedicellate or sessile, epigynous,
beautifully coloured.
Calyx Sepals 3, polysepalous, herbaceous or more or less green to purple coloured; persistent, sepals
persist even in fruit.
Corolla Petals 3, gamopetalous or only basally connate, long one of the petals is usually smaller
than the others; imbricate.
Androecium Highly modified and form the showy part of the flower; stamens usually 2–5, or some-
times 6 or even 4, all briefly basally connate and petaloid; only one stamen is fertile and bears a
single fertile anther cell on its edge, the other stamens are represented by infertile staminodes; one
of the staminodes is reflexed and is called labellum or lip; the other 2 staminodes, if present, are
called wings; 4th staminode, if present, is located behind the fertile stamen; in some species, label-
lum is the only staminode present.
Gynoecium Tricarpellary, syncarpous, trilocular; many anatropous ovules in each locule arranged
in 2 rows; axile placentation; ovary inferior, green and warty; style one, long and petaloid; stigma
one, flat.
Fruit Capsule, warty, usually crowned by persistent sepals (Fig. 22.7); dehiscing by collapse of the
warty pericarp.
Seeds Small, numerous, subglobose, contain very hard endosperm and straight embryo.
General Floral Fromula Br, ≈, , K3, C (3), A1+4 staminodes, G (3).

• Cannaceae is of economic importance because of ornamental value of several species of
Canna and number of their hybrids, e.g. C. indica, C. orchioides, C. generalis, C. orientalis,
etc.
• Edible starch is obtained from the tubers of Queensland Arrowroot (Canna edulis).
484 Plant Taxonomy
staminodes
stigma
style
anther
flower
stamen
petal
labellum
inflorescence
C
A Stamen
B sepal (Upper Part)
ovary
pedicel
A Single Flower
stigma
ovary
leaf style
ovules D
E
Style and Stigma
A
L.S. Inferior Ovary
A Flowering Branch
persistent
sepals
fruit
locules fruit
ovules bracts
F
T.S. Ovary H
A Dehiscing Fruit
G
Fruit
Fig. 22.7 Canna indica L.

• The rhizomes of C. bidentata are sometimes considered as emergency foods.

• The roots of C. indica are diphoretic, diuretic and are given in fevers and dropsy.

Bentham and Hooker treated Canna and its species under Scitamineae of series Epigynae of
Monocotyledons. Engler and Prantl, however, split Scitamineae into 4 independent families, viz.
Musaceae, Cannaceae, Zingiberaceae and Marantaceae. Lymon Benson, however, placed all these
4 families under order Musales, whereas Hutchinson placed all these 4 under order Zingiberales.
Taxonomists treated all these 4 families together because they are all characterised by characters
like (i) presence of 1 or 5 functional stamens, (ii) zygomorphic flowers, (iii) leaves distichous or in
spirals, (iv) inferior ovary, and (v) endospermic seeds.
22.7 MUSACEAE BANANA FAMILY

Monocotyledons, Epigynae, Musaceae (syn. Scitamineae of Bentham and Hooker).

Perennial giant herbs appearing like trees; leaves large, forming crown at the apex of stout unbranched
stem; flowers zygomorphic; inflorescence spadix covered by a spathe; ovary inferior; fruit berry.

A small family of 6 genera (Musa, Ensete, Orchidantha, Ravenala, Heliconia and Strelitzia) and
130 species distributed only in tropical regions. Professor A.D.J. Meeuse (H. de Vries Laboratories,
Amsterdam), while commenting on author’s “A Manual of Practical Botany Vol. II ” in 1975, stated
about the number of genera in Musaceae that “. . . some phanerogamists recognize only Musa and
Ensete (or only Musa if Ensete is not segregated from it) whereas other botanists include the
Heliconiaceae, Lowiaceae and Strelitziaceae in the Musaceae which accounts for their mentioning
5 to 6 genera”1 in this family. Banana (Musa paradisiaca subsp. sapientum) is the most common
plant of the family, found also in India.

General Habit Large perennial herbs, often tree-like in appearance; containing unbranched, tall,
aerial pseudo-stems formed by the leaf sheaths; plants sometimes attain a height of 10 to 15 metres;
plants persist by underground rhizome.
Leaf Very large (up to 2 metres in length), alternate, sometimes distichous or spirally arranged;
oval or oblong with a stout midrib; entire; numerous parallel veins extending up to the margins;
rolled in bud.
1
Personal communication with the author in 1975.
486 Plant Taxonomy
Inflorescence Flowers arranged in racemes and subtended by coloured bracts or spathes; usually
single terminal inflorescence develops from the rhizome and comes out at the top of pseudo-stem
(Fig. 22.8).
Flower Mostly unisexual, plants monoecious having male flowers within the upper bracts of inflo-
rescence and the female flowers within the lower bracts; zygomorphic; trimerous and epigynous;
flowers sometimes bisexual; in Heliconia and Ravenala the bracts are 2-ranked; in Orchidantha,
the flowers are orchid like.
Perianth Tepals 6, arranged in 2 whorls of 3 each; petaloid; unequal-sized; one posterior tepal of
inner whorl is free and boat shaped; remaining 5 tepals (i.e. 2 of inner whorl and 3 tepals of outer
whorl) are united to form a tubular body.
Androecium Stamens 6, arranged in 2 whorls of 3 each; of these 6 stamens 5 are fertile and
antiphilous while 1 posterior stamen of the inner whorl is represented by a staminode (Fig. 22.9); in
Ravenala and sometimes in Ensete all the 6 stamens are fertile; filaments long; anthers dithecous,
basifixed or adnate; introrse; dehiscing by vertical slits; present only in male or bisexual flowers.
Gynoecium Tricarpellary, syncarpous; ovary inferior, trilocular; with one to numerous anatropous
ovules in each locule; axile placentation; style filiform; stigma capitate.
Fruit and Seeds Fruit an elongated berry (Musa), or trilocular capsule (Ravenala, Strelitzia), or
schizocarp splitting into mericarps (Heliconia). Seeds often arillate, with straight embryo in mealy
perisperm.
General Floral Formula (Bisexual Flower) Br, , , P(5)+1, A3+2+1 staminode, G (3).

• Musa paradisiaca L. sub sp. sapientum Schum. (banana or Kela) is one of the most famous
tropical fruits, eaten throughout the world. Over 14 species of Musa occur in India. Green
bananas are used as vegetable, and dried banana powder is a good baby food used in the
manufacture of biscuits, chocolate, etc. Starch is prepared from banana stem. Fibre obtained
from the sheathing leaf bases of M. chinensis is used for cordage. M. textilis also provides
a useful fibre known as Manila hemp.
• Ravenala madagascariensis (Traveller’s Tree), Strelitzia reginae and species of Heliconia
and Orchidantha are most favoured for decorative purposes.
• Ensete ventricosa inflorescences are used as vegetable.

Some taxonomists include only Musa and Ensete in Musaceae, Heliconia in Heliconiaceae,
Orchidantha in Lowiaceae, and Strelitzia and Ravenala in Strelitziaceae. Other phanerogamists divide
Musaceae into 3 subfamilies viz. Lowioideae (Orchidantha), Musoideae (Musa) and Strelitzioideae
stamens
locule
stigma stamen
placenta
ovule
T.S. Ovary
Male Flower Bisexual Flower
leaves
Female Flower
stamen
fruits
stigma flowers
style spathe
tepal
ovary
flowers
L.S. Bisexual Flower
Inflorescence (spathe opened) Flowering plant
Fig. 22.8 Musa paradisiaca L.
(Heliconia, Ravenala and Strelitzia). Bentham and Hooker

included only Musa and Ensete in tribe Museae of family
Scitamineae.
Musaceae is closely related to other families of
Scitaminales, and originated from Liliaceous stock. Engler
believed Musaceae to be the ancestral stock of Orchidaceae.
Takhtajan (1969) treated Musaceae, Heliconiaceae, Lowiaceae
and Strelitziaceae under order Zingiberales along with 4 other
families and believed them to be derived from Liliales.
Fig. 22.9 Floral diagram of
bisexual flower of Musa paradisiaca.
488 Plant Taxonomy
22.8 ZINGIBERACEAE GINGER FAMILY

Monocotyledons, Epigynae, Zingiberaceae.

Aromatic herbs; ligule present at the top of leaf sheath; perianth differentiated into calyx and corolla;
only single fertile stamen; staminodes petaloid.

Amomum, Curcuma, Elettaria, Zingiber.

A family of 49 genera and over 1000 species, Zingiberaceae are distributed in tropical regions,
chiefly in Indomalaysia. About 17 genera and over 115 species have been reported from India, chiefly
in Western Ghats and Eastern Himalayas. Some largely represented genera along with the number
of their worldover reported species are Alpinia (250), Amomum (150), Costus (150), Zingiber (90),
Kaempferia (70), Curcuma (60), Hedychium (50), Globba (50) and Elettaria (7).

General Habit Perennial, often aromatic herbs; usually with sympodial tuberous (Curcuma) or
horizontal (Zingiber) rhizomes; rarely with fibrous roots (Cautleya); aerial stem, if present, is short;
sometimes an apparent stem is formed by the rolled-up leaf-sheaths.
Leaf Alternate, sessile or petiolate, distichous or spirally arranged, with sheathing leaf bases; ligu-
late, ligule present at the junction of blade with petiole or sheath; leaf blade large, linear to elliptical,
with closely parallel-pinnate venation.
Inflorescence Variable from a compact spike (Alpinia, Zingiber), raceme (Curcuma), panicle
(Globba), or flowers solitary; cymose in some species of Globba; each flower or group of flowers
subtended by a large bract; in several genera the inflorescence develops on leafless scapes.
Flower Bracteate, pedicellate or sessile, bisexual, zygomorphic, only rarely actinomorphic, trim-
erous, epigynous, large, brightly coloured, usually aromatic or fragrant; perianth differentiated into
calyx and corolla.
Calyx 3 united sepals forming a tubular or spathiform, green or herbaceous calyx; the odd sepal
is anterior; calyx tube is very short in Globba and splitted in Roscoea.
Corolla 3 unequal-lobed petals, free or more or less united into a tubular corolla with the posterior
lobe usually the largest; delicate and usually showy.
Androecium Basically 6 stamens, in 2 whorls of 3 each; 2 lateral stamens of the outer whorl modify
into staminodes while the 3rd anterior stamen of this whorl is usually absent or highly suppressed; 1
median or posterior stamen of the inner whorl is fertile and 2 lateral stamens of this whorl unite to
anther
filament enclosing fertile stamen

style petal
corolla lobe sepal
labellum
staminode
corolla tube
staminode
of
calyx tube outer whorl
suppressed
labellum stamen of
bract outer whorl
inferior ovary
Fig. 22.10 A flower of Hedychium. Fig. 22.11 Floral diagram of Curcuma longa.
form a labellum (Figs. 22.10, 22.11); fertile stamen is 2-celled, usually epipetalous, with its filament
usually slender and deeply grooved.
Gynoecium Tricarpellary, syncarpous; ovary inferior, usually trilocular with numerous anatropous
or semianatropous ovules on axile placentation, or unilocular with 3 parietal placentae (Globba);
style usually filiform and more or less enveloped in the groove of filament of fertile stamen; stigma
protruding beyond the anther lobes; usually epigynous nectar-secreting gland present.
Fruit and Seeds Fruit usually a 3-valved, loculicidal capsule, or rarely fleshy, indehiscent, berry-like.
Seeds often arillate, and each with straight embryo and abundant endosperm.
General Floral Formula Br, , , K(3), C (3) or 3, A1, G (3).

Economically, the family is important as a source of some spices and condiments, fragrant oils,
and ornamentals.
• Spices and Condiments Seeds of Amomum cardamon (cardamon or Choti Elayachi) and
Elettaria cardamomum (true cardamon or Elayachi), roots of Alpinia officinarum (gangal
root), and rhizomes of A. gangala (Siamese ginger), Curcuma longa (turmeric or Haldi) and
Zingiber officinale (ginger or Adrak) are used as spices, condiments and flavouring agents.
Ginger oil, obtained from the rhizome of Zingiber officinale, is used in perfumery and for
medicinal purposes. Turmeric is also used for medicinal purposes.
490 Plant Taxonomy
• Abir Abir, the famous scented powder, is prepared from Hedychium spicatum (Kafur-
Kachri).
• Zedoary The product ‘Zedoary’, used as a tonic and in perfumery, is obtained from the
tubers of Curcuma zedoaria.
• Cultivated Ornamentals Some of the ornamental genera cultivated in greenhouses and gar-
dens for decorative purposes are Alpinia (shell ginger), Brachychilum, Cardamon, Cautleya,
Costus, Globba, Hedychium, Kaempferia and Roscoea.

Treated only as a tribe Zingiberae of the family Scitamineae by Bentham and Hooker, Zingiberaceae
has been treated as an independent family of the order Zingiberales by Hutchinson (1959); Takhtajan
(1969), Cronquist (1981) and Thorne (1983). It is divided into 2 subfamilies viz. Zingiberoideae (plants
aromatic, with oil cells, leaves 2-ranked) and Costoideae (plants without oil cells, leaves spirally
arranged). Willis treated Costus and a few related genera in an independent family Costaceae.
Zingiberaceae is closely related to Musaceae in habit, zygomorphic flowers and inferior ovary.
However, due to the reduction in androecium, Zingiberaceae has been considered to be more
advanced.
22.9 LILIACEAE LILY FAMILY

Monocotyledons, Coronarieae, Liliaceae.

Perennial herbs often with bulbs, corms, or rhizomes; leaves usually radical; perianth showy, usually
of 6 tepals, arranged in 2 whorls of 3 each; stamens 6; ovary superior.

Allium, Aloe, Asparagus, Asphodelus, Colchicum, Dracaena, Fritillaria, Lilium, Ruscus, Sansevieria,
Smilax and Yucca.

A family of about 250 genera and 3700 species showing cosmopolitan distribution. About 35 genera
and over 195 species have been reported from India. Some of the largely represented genera along
with the number of their worldover reported species are Allium (450), Smilax (350), Aloe (300),
Asparagus (300), Dracaena (150), Ornithogalum (150), Haworthia (150), Tulipa (100), Fritillaria
(85), Lilium (80), Colchicum (65), Sansevieria (60), Yucca (40), Asphodelus (12) and Ruscus (7).

General Habit Mostly perennial herbs with sympodial bulbs (Allium cepa) , rhizomes (Paris quadri-
folia), corms (Colchicum), or bulbils (Lilium); some are woody shrubs (Dracaena, Yucca), or trees
(Xanthorrhoea); a few are succulents (Aloe, Gasteria), and climbers (Gloriosa, Smilax); in some, the
ultimate branches are modified into phylloclades (Ruscus) or cladodes (Asparagus) and the leaves
are reduced to scales.
Leaf Basal or cauline, simple; exstipulate but stipulate in Smilax where the stipules modify into
tendrils; usually alternate, less commonly whorled (Paris), and rarely opposite (Gloriosa, Scolyopus);
sometimes fleshy or spiny margined (Aloe), acicular, long, fleshy and hollow in Asphodelus
(Fig. 22.12); reduced to scales in Ruscus and Asparagus; usually parallel venation but in Smilax
and Paris the venation is reticulate.
Inflorescence Usually racemose raceme (Asphodelus), sometimes in spikes (Aloe), terminal panicle
(Yucca), or monochasial cyme in Hemerocallis, or cymose in apparent umbels in Allium, Agapanthus;
flowers solitary axillary (Gloriosa), or solitary terminal (Tulipa).
Flower Bracteate, usually ebracteolate except Dianella and Lilium; bisexual, actinomorphic, trim-
erous, hypogynous; rarely unisexual (Smilax zeylanica, Fig. 22.13 A, B; Ruscus) and such species
are usually dioecious; rarely tetramerous (Maianthemum, Paris); slightly zygomorphic (Lilium,
Haworthia).
ovary wall
locule flower
anther
ovule lobe
placenta
filament
T.S. Ovary
A Stamen inflorescence
leaves
roots
Floral Diagram
A Flowering Plant
Fig. 22.12 Asphodelus tenuifolius L.

492 Plant Taxonomy
A B
Fig. 22.13A, B Smilax zeylanica: Floral diagrams—A: Male flower; B: Female flower.
Perianth 6 tepals arranged in 2 whorls of 3 each, free or rarely united (Aloe); usually petaloid or
sometimes sepaloid; odd tepal of the outer whorl is anterior in position; valvate or imbricate; more
than 6 tepals in Paris quadrifolia.
Androecium Usually 6 stamens arranged in 2 whorls of 3 each; rarely the stamens are 3 (Ruscus),
4 (Maianthemum) or up to 12; polyandrous; opposite the tepals; sometimes epiphyllous; filaments
distinct or connate; anthers dithecous, basifixed or versatile, extrorse or introrse, dehiscing usually
by vertical slits and sometimes by terminal pores; rarely synandrous (Ruscus).
Gynoecium Tricarpellary, syncarpous, the odd carpel usually anterior; ovary superior, trilocular,
with 2 rows of numerous anatropous ovules, axile placentation; rarely unilocular with parietal pla-
centation; style usually 1; stigmas 1 or 3; rarely the ovary is inferior (Haemodorum); usually the
nectar-secreting septal glands are present in the ovary.
Fruit and Seeds Fruit usually a septicidal or loculicidal capsule or a berry Asparagus, Smilax). Seeds
with curved or straight embryo and abundant endosperm.
Pollination Flowers are usually insect-pollinated. Insects are attracted usually because of gaudy
and scented perianth lobes and the nectar secreted in the septal glands. In Paris, the insects are
attracted because of the foetid smell and dark purple colour of the floral parts. Bright colours of
tepals in Tulipa also attract insects. Snails which come to eat the fleshy tepals of Rhodea bring
about the cross pollination.
In Yucca the cross-pollination is carried out by a special moth, Pronuba yuccasella. Fully
expanded flowers emit perfumes and are visited by the female moth, especially during nights. She
collects a lot of pollen grains from one flower and visits another flower. Life-history of this moth
is intimately associated with the pollination mechanism in Yucca.
General Floral Formula Br, Ebrl, ≈, , P3+3, A3+3, G (3).

Family is important from economic point of view in several ways, of which some are
undermentioned:
• Edible Plants (i) Bulbs of Allium cepa (onion, Piyaz), used throughout the world as veg-
etable, possess stimulative, diuretic, expectorant and bactericidal properties, (ii) Bulbs of
Allium sativum (garlic, Lahsun), used as condiment like onion, are also good for heart,
(iii) Fleshy shoots of Asparagus officinalis and tuberous roots of A. racemosus are used as
vegetable.
• Medicinal Plants (i) Aloe barbadense (Ghikanwar) leaves are the source of resinuous drug,
used as a purgative, (ii) Aloe vera (Aloe) leaves provide mucilaginous liquid, used in piles
and inflammations, (iii) Medicated oil prepared from the roots of Asparagus racemosus is
used for nervous and rheumatic complaints and also in skin diseases, (iv) Colchicum luteum
roots are used in the treatment of gout and rheumatism, (v) Fritillaria roylei bulbs are used
as expectorant and antipyretic, (vi) Gloriosa superba tubers prove helpful in promoting labour
pains in women, (vii) Hemerocallis fulva flowers are blood purifier and given to women
during child birth, (viii) Iphigenia indica corms are used in stomach pains and headache,
(ix) Paris polyphylla rhizomes are used as anthelmintic, (x) Scilla hyacinthiana bulbs are
used as heart stimulant, (xi) Smilax glabra and S. ovatifolia roots are used in the treatment
of venereal diseases, (xii) Urginea indica bulbs are heart stimulant and also used in skin
diseases and rheumatism, (xiii) Drug obtained from the roots of Veratrum viridae is used
in hypertension.
• Resins Resins are obtained from the stems of Dracaena and Xanthorrhoea.
• Fibre Plants Excellent fibres, used for cordage, fishing nets, mattings, twines etc., are
obtained from Phormium tenax (New Zealand Flax), Sansevieria roxburghiana, Smilax
glabra and Yucca filamentosa.
• Insecticides and Raticides Bulbs of Urginea indica are used for killing rats while that of
Veratrum album are used as insecticides.
• Polyploidy Colchicine, an alkaloid obtained from Colchicum luteum, is used to induce
polyploidy.
• Ornamental Plants Some of the well known garden ornamentals of Liliaceae are
Agapanthus africanus (African lily), Asparagus plumosus, Dracaena, Fritillaria tenella,
Gloriosa superba (Malabar Glory Lily), Hemerocallis fulva (Orange Day Lily), Lilium can-
didum (Lily), L. giganteum, Ruscus aculeatus (Butchers broom), Tulipa suaveolens (Tulip),
Yucca aloifolia and Y. gloriosa.

Discussed under the series Coronarieae by Bentham and Hooker, Liliaceae was treated under
Liliiflorae by Engler and Prantl (1931), and Liliales by Hutchinson (1959), Takhtajan (1969), Cronquist
(1981) and Thorne (1983). Cronquist merged Amaryllidaceae under Liliaceae. Some taxonomists
494 Plant Taxonomy
include certain genera of Liliaceae under independent families, e.g. Aloe under Aloeaceae and Smilax
under Smilacaceae. Bentham and Hooker divided Liliaceae into 20 tribes whereas Engler and Prantl
divided it into 12 subfamilies.
Family Liliaceae represents most primitive basic monocotyledonous stock from which the other
families of monocots have originated. According to Takhtajan (1969), order Liliales, together with
orders Alismales and Triuridales “have more probably a common origin from a hypothetical extinct
group with endospermous seeds and 2-celled pollen grains, as in the Liliales”, and subfamily
Melanthioideae “of the family Liliaceae is nearest to the ancestral type”.

Asphodelus tenuifolius Cav. (vern. Piazi, Fig. 22.12)
Habit: An annual weed with fibrous roots, and condensed, underground and reduced stem.
Leaf: Radical, simple, exstipulate, sessile, with sheathing leaf bases; long, acicular, acute, fleshy,
hollow. Inflorescence: Racemose raceme with flowers on long scape or peduncle. Flower: Bracteate,
pedicellate, bisexual, actinomorphic, trimerous. Perianth: 6 tepals arranged in 2 whorls of 3 each,
free, petaloid, valvate. Androecium: 6 stamens arranged in 2 whorls of 3 each, polyandrous, epiphyl-
lous, present opposite the tepals. Gynoecium: Tricarpellary, syncarpous, superior, trilocular, 2 ovules
in each locule; axile placentation; style long; stigma trifid. Fruit: Capsule.
Floral Formula: Br, ≈, , P3+3, A3+3 , G (3).
22.10 COMMELINACEAE SPIDERWORT FAMILY

Monocotyledons, Coronarieae, Commelinaceae.

Herbs with succulent stem, and leaves with tubular sheath; floral parts in 3s; perianth differentiated
into calyx and corolla; ovary superior; fruit capsule.

Aneilema, Commelina, Murdannia, Streptolirion and Tradescantia.

A family of about 50 genera and 700 species, Commelinaceae are mostly tropical and subtropical
in distribution. About 11 genera and over 75 species have been reported from India, chiefly from
eastern Himalayas and parts of southern and western India. Some larger genera along with the total
number of their reported species and common names of some of them are Commelina (200, day-
flower), Aneilema (100), Tradescantia (65, spiderwort), Cyanotis (30), Dichorisandra (30), Zebrina
(4, wandering Jew) and Rhoeo (1, Moses-in-the bulrushes).

General Habit Annual or perennial herbs with fibrous or tuber-like roots, jointed and more or less
succulent stems; occasionally twining (Streptolirion); stem either with purplish tinge (Commelina)
or deep violet (Tradescantia).
Leaf Simple, alternate, sheathing at the base, flat or trough-like, linear or ovate or lanceolate; entire
margins; parallel-veined.
Inflorescence Usually an umbellate cyme subtended by two large spathaceous bracts (Tradescantia),
thyrsoid panicles (Floscopa), or flowers solitary.
Flower Pedicellate; usually actinomorphic but zygomorphic in Commelina; bisexual; usually sub-
tended by boat-shaped spathe or foliaceous bract; trimerous, hypogynous, commonly blue; in
Commelina some flowers are also subterranean and cleistogamous.
Perianth Usually biseriate i.e. differentiated into calyx and corolla; calyx is represented by an outer
whorl of 3 sepal-like green tepals which are usually persistent, free and imbricate, and only rarely
connate; corolla is represented by an inner whorl of 3 petal-like, usually ephemeral, coloured, equal or
unequal-sized tepals which are also usually free and imbricate, and only rarely united (Cyanotis).
Androecium Typically 6 stamens in 3 + 3 arrangement, distinct or rarely fused, frequently some
reduced to staminodes; there are 2 fertile stamens and 4 staminodes in Aneilema, 3 fertile stamens
and 3 staminodes in Commelina (Fig. 22.14), 5 fertile stamens and 1 staminode in Floscopa, and all
the 6 fertile stamens and no staminode in Tradescantia (Fig. 22.15); filaments often hairy or bearded;
anthers 2-celled (parallel or divergent), dehiscence longitudinal or poricidal (Dichorisandra).
Gynoecium Tricarpellary, syncarpous; ovary superior, trilocular, with 1 to few orthotropous ovules
in each locule; axile placentation; style 1; stigma 1, capitate or trifid.
Fruit and Seeds Fruit a loculicidal capsule, rarely fleshy and indehiscent. Seeds often arillate, 1
to few in each locule, with small embryo and copious endosperm; seed tissue contains calcium
oxalate.
Pollination and Dispersal Flowers in Tradescantia and several other genera are protandrous and
adapted for insect pollination. As the flowers fade, the petals become pulpy and their surface gets
covered with a layer of liquid which attracts insects. Fruits are dispersed by animals and birds.
General Floral Formula Br, ≈ or , , K3, C3, A3+3 or 3+3 staminodes or 2+4 staminodes or 5+1 staminode, G (3).

• Except for some ornamental plants, the family is of little importance. The ornamental spe-
cies include Commelina suffruticosa, Cyanotis cristata, Dichorisandra thyrsiflora, Rhoeo
discolor, Tradescantia virginiana (spiderwort) and Zebrina pendula (wandering Jew).
• Some minor aspects of utility include: (i) the roots of Aneilema scapiflorum are used in piles
and asthma, (ii) the rhizomes of Commelina benghalensis are used as vegetable, (iii) the
roots of C. obliqua are used as an antidote to snake poison, (iv) the stem juice of Floscopa
scandens is put in sore eyes, (v) plants of several species of Aneilema, Murdannia and
Commelina are used in leprosy and leucoderma, (vi) roots of Cyanotis are used to expel
worms in cattles.
496 Plant Taxonomy
fertile stamen
stigma
staminode flower
style
ovary
staminode
L.S. Flower
sepal
leaf
petal
stamen
An Opened Flower
stem
Floral Diagram Flowering Plant
Fig. 22.14 Commelina benghalensis L.

(Affinities)
The family is included under series Coronarieae by
Bentham and Hooker, order Farinosae by Engler,
and under order Commelinales by most of the
other workers including Hutchinson, Takhtajan,
Cronquist, Thorne, etc. Commelinaceae are usually
divided into 2 tribes viz. Tradescantieae (flowers
usually actinomorphic) and Commelineae (flowers
usually zygomorphic).
Hutchinson (1959) derived order Commelinales
from 2 most primitive orders viz. Butomales and
Alismatales, i.e. from Helobiae. However, Takhtajan
(1969) opined that Commelinales “probably has
Fig. 22.15 Floral diagram of Tradescantia.
a common origin with the Bromeliales” from
Liliales.

Commelina benghalensis L. (Fig. 22.14)
Habit: Annual weed with semi-erect or creeping stem having nodes and internodes. Leaf: Simple,
exstipulate, sessile with sheathing leaf base, ovate to oblong-lanceolate, multicostate parallel venation;
greenish with purplish tinge. Inflorescence: Cymes surrounded by 1–3 spathes or bracts. Flower:
Bracteate, pedicellate, bisexual, zygomorphic, hypogynous, trimerous, blue coloured; some flow-
ers subterranean and cleistogamous. Perianth: Biseriate; outer whorl of 3 tepals represents calyx,
polysepalous, valvate or imbricate; inner whorl of 3 represents corolla, polypetalous, 1 petal larger;
imbricate. Androecium: 6 stamens of which 3 are fertile and 3 are sterile staminodes; antiphyllous,
light blue filaments, anthers dithecous, discrete and divaricate. Gynoecium: Typical as discussed for
the family, except that 1 locule contains 1 ovule and the other 2 locules contain 2 ovules each.
Floral Formula: Br, , , P3+3, A3+3 staminodes, G (3).
22.11 JUNCACEAE RUSH FAMILY

Monocotyledons, Calycineae, Juncaceae.

Grass-like or rush-like, annual or perennial, tufted herbs; leaves basal, tufted, linear, grass-like,
sheathing basally or reduced to basal sheath; perianth usually sepalloid, segments 6, in 2 whorls of
3 each; stamens 6, generally in 2 whorls of 3 each; ovary tricarpellary, syncarpous, superior; fruit
capsule.

Juncus, Luzula.

Juncaceae includes about 400 species of only 8 genera (Juncus, Luzula, Rostkovia, Marsippospermum,
Andesia, Oxychloe, Distichia and Prionium), of which only 2 (Juncus and Luzula) are found in India.
The two Indian genera are cosmopolitan in their distribution. Juncaceae members are distributed
mainly in temperate and arctic regions and also in tropical mountains, specially in moist cool places.
The chief genera are Juncus (300, rush; Fig. 22.16) and Luzula (80, wood rush). Rushes often provide
stepping stones for crossing a wet meadow.

General Habit Annual or perennial tufted herbs, only rarely shrub-like (Prionium), with their stems
usually leafy only at the base; these plants of damp and cool places usually contain a creeping
sympodial rhizome, of which one joint usually appear above ground each year and develop into a
leafy shoot.
498 Plant Taxonomy
Leaves Basal, narrow, flat or cylindrical; grass like, sessile; linear or filiform; sheathing at the base
or reduced to only a sheath; tufted; exstipulate; mostly hairy and parallel-veined.
Inflorescence Flowers usually a crowded mass of cymes arranged in a panicle, corymb or head;
rarely flowers solitary.
Flowers Usually bracteate, pedicellate, sessile, bisexual, actinomorphic, trimerous, small and wind-
pollinated; if unisexual, then the plants are dioecious (e.g. Oxychloe); hypogynous.
Perianth Segments 6, arranged in 2 whorls of 3 each; polyphyllous; usually sepalloid or green,
rarely coloured; scale-like or glumaceous or coriaceous.
Androecium Stamens 6, arranged in 2 whorls of 3 each, or inner whorl absent; antiphyllous (opposite
the perianth lobes); polyandrous; anthers dithecous, basifixed, introrse; dehiscing by vertical slits;
pollen in tetrads.
Gynoecium Tricarpellary, syncarpous, superior; unilocular, with 3 parietal placentae (e.g. Luzula),
or trilocular with one to three or many anatropous ovules in each locule showing axile placentation;
styles 1–3, simple; stigmas 3, brush-like.
Fruit A 1–3 loculicidal capsule.
Seeds Small, with straight embryo and starchy endosperm.
General Floral Formula Br, ≈, , P3+3, A3+3, G (3).

Juncaceae are of little economic importance. In some countries, various species of Juncus (e.g.
J. effusus) are woven into mats, hats, chair seats, baskets, etc. Pith of some of the rushes is used
for candlewicks. Few rushes are grown for ornamental purposes in locations adjacent to aquatic
habitats, and also used horticulturally for waterside planting. Juncus squarrosus is used as fod-
der for sheeps. Some species of Luzula (e.g. L. pilosa, L. campestris) are used in kidney trouble.
Paper is made from fibres obtained from some species of Luzula.

Juncaceae has been included under series Calycineae by Bentham and Hooker along with two more
families, Palmaceae and Flagellariaceae. Engler and Prantl, however, discussed Juncaceae under order
Liliales. Hutchinson included Juncaceae under order Juncales of division Glumiflorae.
Due to the presence of typical trimerous flowers with superior ovary, both Junaceae and Liliaceae
are said to be quite close to each other, and Juncaceae is considered to be derived from Liliaceae
by reduction. Anatomical studies of two families, however, do not support this view. Due to the
similarities in the distribution of vessels and type of stomata, Juncaceae resemble with families like
Cyperaceae, Restionaceae and Gramineae. Morphology and pollen development of Juncaceae and
Cyperaceae are quite similar and these bring the two families together.
inflorescence
stigma
style
superior ovary
stamen
outer perianth
segment
B flowering
inner perianth
stem
segment (folded back)
A flower of Juncus
tuft of
leaves
locule
C
ovules
A Whole Plant of
perianth Luzula campestris
locule
F ovules
A L.S. Superior Ovary

Inflorescence of G
Juncus inflexus T.S. Ovary
stigmas (3)
ovary stamen
style
stamens
perianth
lobes
perianth
lobe
ovary
D E
A Bisexual Flower Parts of L.S. of a Flower
Fig. 22.16 A, B, Juncus inflexus; C–G, Various floral parts of Luzula campestris.
500 Plant Taxonomy
22.12 PALMAE OR ARECACEAE PALM FAMILY

Monocotyledons, Calycinae, Palmae or Arecaceae.

Shrubs or trees with arborescent stem having prominent scars of leaf bases and crown of large
fan-shaped or pinnately compound leaves; inflorescence spadix or paniculate, often with spathes;
flowers-small, unisexual or bisexual, actinomorphic, trimerous.

Areca, Borassus, Calamus, Cocos, Nypa, Phoenix.

A family of 217 genera and over 3000 species distributed widely in tropical and subtropical regions
of the world, and a few also in warm temperate regions. About 28 genera and over 95 species have
been reported from India. Chief genera along with the number of their total reported species and
the common names of some of them are Calamus (300, cane palm), Chamaedorea (130), Areca (54,
betel-nut palm), Coccothrinax (50, biscayne palm), Copernicia (30, wax palm), Raphia (30, wine
palm), Sabal (25, cabbage palm), Phoenix (17, date palm), Roystonea (17, royal palm), Metroxylon
(15, sago palm), Phytelephas (15, ivory-nut palm), Caryota (12, fish-tail palm), Elaeis (2, oil palm)
and Cocos (1, coconut palm).

General Habit Woody shrubs, trees, or sometimes vines (Calamus, Desmoncus); with usually
unbranched, slender to stout stem; short to over 30 metres tall; and with leaves often forming a
terminal cluster in the arborescent species.
Root Fibrous and adventitious roots arising from the base of stem; in Iriartea prop roots develop.
Stem Variable in different forms, such as (i) very short with leaves appear to arise from the ground
(e.g. Nypa), (ii) thin and slender with long internodes (e.g. Calamus), or (ii) tall, stout, pillar like,
covered by persistent leaf bases and containing a terminal cluster of leaves (e.g. Cocos, Phoenix);
stem is rarely branched.
Leaf Leaves are palmate (fan palms) or pinnate (feather palms), and rarely simple; usually large
with petiole base often sheathing the stem; usually in a terminal cluster; usually alternate in climb-
ing species (Calamus); petiole is large, strong, smooth or spiny; ochreate stipule present in Calamus
while leaves are ligulate in Thrinax; parallel venation.
Inflorescence Large, much branched, paniculate (Daemonorops) or spadix (Phoenix), often reach-
ing up to 1 metre, and covered by one or more boat-shaped woody spathes or bracts; interfoliar i.e.
arise amongst the leaves(e.g., Borassus) or intrafoliar i.e. arise below the leaves (e.g. Areca); plants
monoecious or dioecious or flowers bisexual.
Flower Ebracteate, sessile, actinomorphic; usually unisexual and monoecious but sometimes dioe-
cious (Phoenix), rarely bisexual (Livistona); trimerous, hypogynous, in Borassus, the male flowers
are smaller than female flowers.
Perianth 6 free or united tepals in 2 whorls of 3 each; tough, leathery and usually persistent;
imbricate, valvate or twisted in the bud; in Phoenix the tepals of outer whorl are united and valvate
while that of inner whorl are free and twisted (Fig. 22.17); usually the tepals of outer whorl are
smaller than of inner whorl.
Androecium Present only in staminate or bisexual flowers; usually 6 stamens arranged in 2 whorls
of 3 each (Phoenix); positioned against tepals; filaments short, distinct; anthers dithecous, basifixed
anther
lobe
Female
Inflorescence Stamen
Male
Inflorescence
stigma anther
lobe
tepals tepals
(inner) (inner)
tepals tepals
(outer) (outer)
Female Flower leaflet L.S. Male Flower
Leaf
Floral Diagram (Female flower) Floral Diagram (male flower)
Fig. 22.17 Phoenix sylvestris (L.) Roxb.

502 Plant Taxonomy
or dorsifixed, introrse, dehiscence by vertical slits; stamens are only 3 in Nypa and numerous in
Caryota and Phytelephas.
Staminodes and Pistillodes A pistillode is usually present in the centre of each male flower.
Staminodes may (Phytelephas) or may not be present in the female flowers.
Gynoecium Present only in pistillate or bisexual flowers; tricarpellary, syncarpous; or carpels are
partly (Nypa) or completely (Phoenix) free; ovary superior, 3-locular with a single anatropous ovule
in each locule; axile placentation; in Phoenix the placentation is basal; sometimes the ovary is uni-
locular and the placentation is parietal; style extremely short or absent; stigmas 3, sessile.
Fruit and Seeds Fruit berry with fleshy exocarp (Phoenix) or drupe with fibrous epicarp (Cocos).
Seeds with small embryo and abundant endosperm.
Pollination and Dispersal Pollination in palms is usually anemophilous, and the flowers are
protandrous. Some palms (Sabal), however, are insect pollinated. Dispersal of fruit is either by
animals or by water (Cocos).
(a) Male flower: Ebr, ≈, , P(3)+3 or 3+3, A3+3, G0 or pistillode.
(b) Female Flower: Ebr, ≈, , P(3)+3 or 3+3, A0 or staminodes, G (3) or 3.
(c) Bisexual Flower: Ebr, ≈, , P(3)+3 or 3+3, A3+3, G (3) or 3.

Palm family, supposed to be second in importance to Poaceae, is used for food, oils, shelter,
coconuts, copra, dates, clothing, etc. As many as 801 uses of only Borassus flabellifer have been
mentioned in one ancient Tamil song. Cocos nucifera (coconut palm or Nariyal) is another such
plant of this family whose almost every part of the entire plant body is utilized in some ways or
other, i.e. its (i) root decoction is used as mouthwash, (ii) stem is used for pillars and as a fuel,
(iii) leaves are utilized for making mats, fans, baskets, etc., (iv) toddy, obtained from the young
inflorescences, provides an intoxicating alcoholic drink on fermentation, (v) endosperm of young
fruits is edible and their milky fluid is a sweet and refreshing drink; (vi) vegetable fat, obtained
from the dried endosperm of mature fruits, is used for cooking and several toilet preparations,
(vii) dried fibrous mesocarp of fruits is used for brushes, ropes, mats, floor coverings, etc. (viii)
woody endocarp of fruits is prepared into several articles of decoration, and (ix) famous coconut
oil is used for anointing the body.
Some of the other similar generalized uses of Palmae are undermentioned:
• Ornamental Palms: Palms are universally known for their majestic look and handsome
appearance. Some of them are Adonidia merilli (Manila palm), Arenga saccharifolia (Gomuti
palm), Caryota urens (wine palm or fish-tail palm), Chamaerops humilus (European
fan palm), Livistona chinensis (Fountain palm or Chinese fan palm), Phoenix rupicola,
Pritchardia pacifica, Roystonea elata (Floridean royal palm), R. obracea, R. regia (Royal
palm or bottle palm), Sabal minor (Bush palmetto) and S. umbraculifera (Cabbage palm).
• Oils: Coconut oil is obtained from the dried ripe endosperm of Cocos nucifera while
the palm oil is obtained from the fleshy mesocarp of the fruits of Elaeis guineensis (oil
palm).
• Edible Products: (i) Seeds of Areca catechu (Supari or betel nut palm) are sliced and
chewed along with “Paan” (Piper betel) by millions of the people, (ii) kernel of fruits and
tender leaves of Arenga pinnata is edible, (iii) young seedlings of Borassus flabellifer are
used as vegetable and its fruits are eaten after roasting, (iv) fruits of Copernicia ceribera and
Hyphaene thebaica are edible, (v) Sago, used as an article of diet, is obtained from the pith
of the stem of Metroxylon rumphii (sago palm or Sabudana), (vi) stem buds, young peduncles
and immature seeds of Nypa fruticans are eaten raw or cooked as vegetable, (vii) ripe fruits
of Phoenix dactylifera (date palm or “Pind Khajoor”) and P. sylvestris (wild date palm or
“Jangli Khajoor”) are eaten throughout the world for their high food value, (viii) fruits of
Sabal palmetto (cabbage palm) are edible, and honey is prepared from its flowers.
• Toddy: A sap or sugary solution, obtained by cutting or tapping the stems, young peduncles
or inflorescence of several palms, is used for manufacturing jaggery, and by fermentation it
yields an intoxicating drink or beverage called “toddy”. This sap is also used for manufac-
turing number of commercially useful products such as ‘Nira’, palm sugar, alcohol, vinegar,
etc. Palms, commonly used for this purpose are Arenga pinnata, A. saccharifera, Borassus
flabellifer, Caryota urens, Metroxylon vinifera, Nypa fruticans, Phoenix dactylifera and P.
sylvestris. The sap of Raphia hookeri is the source of famous ‘Bourdon wine’.
• Hats, Mats and Baskets: These and many other similar articles such as umbrella handles,
walking sticks, hand fans, etc, are prepared from Borassus flabellifer, Nypa fruticans,
Phoenix dactylifera, P. sylvestris, etc.
• Brushes: Brushes are prepared from fibres obtained from the leaf stalks of Arenga pinnata,
Borassus flabellifer and Sabal palmetto.
• Timber: Dense stem wood of several palms (Borassus, Caryota, Cocos, Hyphaene and
Phoenix) is used as timber.
• Cane: Common ‘cane’ or ‘rattan’ of commerce, used for matting chairs, baskets, furniture
frames, walking sticks etc. is obtained from several species of Calamus (Cane or Rattan
palms), such as C. extensus, C. latifolius, C. ovatus, C. rotundus and C. tenuis. Daemonorops
adspersus, D. jenkinsianus and Korthalsia horrida are also utilized for similar purposes.
• Vegetable Ivory: Stony endosperm of Phytelephas macrocarpa is used for making beads
of necklaces and other similar articles as cheap substitute of ivory.
• Wax: Wax, utilized for manufacturing candles, boot polishes, etc. is obtained from
Ceroxylon andicola, Copernicia cerifera, etc.
• Sacred Writing Material: Narrow strips of leaves of Corypha umbraculifera have been used
for several sacred writings in olden days. Leaves of Borassus flabellifer were also used for
several such writings by ancient Hindus.
• Resin: Dragon’s blood, a resinuous exudate obtained from the fruits of Daemonorops ruber,
is used in colouring marbels, varnishes, etc.
504 Plant Taxonomy

Palmae or Arecaceae was variously placed under series Calycinae by Bentham and Hooker, order
Principes by Engler and Prantl, order Palmales by Hutchinson, and order Arecales by Takhtajan,
Cronquist, and Thorne.
Palmae is closely related to Araceae, and it was discussed by Rendle under Spadiciflorae along
with Araceae and Lemnaceae. Hutchinson, however, opined that Palmae is allied to Pandanaceae on
one hand and Agavaceae on the other. He believed that Palmae probably originated from Agavaceae.
Workers like Takhtajan and Cronquist opined that because of the presence of unisexual inconspicuous
flowers developing in spadix in both Palmae and Araceae, the two families are closely related.
22.13 TYPHACEAE CATTAIL FAMILY

Monocotyledons, Nudiflorae, Typhaceae.

Monoecious, large perennial herbs of fresh to brackish open marshes; leaves alternate, erect, long,
linear, in 2-ranks, sessile and with parallel venation; inflorescence a dense spike, fuzzy-brown at
maturity, usually divided into upper part containing male flowers and lower part bearing female
flowers; fruit achene.
22.13.3 Indian Genus

Typha.

Typhaceae is a unigeneric family comprising only 1 genus (Typha, Fig. 22.18) and about 20 spe-
cies distributed in temperate and tropical regions throughout the world. Due to the dense spike-like
inflorescence and long grass-like leaves, this is commonly called water grass or cattail family. Typha
angustata, T. elephantina and T. laxamanni are some common Indian species.

General Habit Erect perennial herbs; rootstocks rhizomatous, rhizome thick and creeping; fibrous
adventitious roots, arise from the nodes of rhizome; the aerial shoot, developing from the axil of
the scaly leaves, terminate into an inflorescence.
Leaves Alternate, long, linear, sessile, mostly basal, developing in 2 ranks; sheathing at the base;
venation parallel; leaves often overtop the inflorescence.
Inflorescence Thick, dense cylindrical spikes, usually divided into two parts: the upper, usually
yellowish part bearing male flowers, and the lower, usually brownish part bearing female flowers;
the two parts usually contiguous.
staminate flowers
male stigma
pistillate flowers flowers
style
hairs
inflorescence ovary
female
flowers
leaves
gynophore
A erect stem E
main axis
A Part of Main Axis With
One Female Flower
rhizome
A Plant With Inflorescence
B
stamens
withering (3)
style
D common
An Inflorescence stalk
A Single Male Flower

fruit
style
male flowers (3) perianth hairs

(reduced)
stamens
G
main axis
A Fruit ovary
gynophore C
F
L.S. Ovary Main Axis with
3 Male Flowers
Fig. 22.18 Typha latifolia L.

506 Plant Taxonomy
Flowers Unisexual, surrounded by slender hairs or more or less forked scales, representing possibly
the reduced perianth; flowers borne on a cylindrical spadix; bracteate, shortly pedicellate, actino-
morphic, hypogynous.
Male Flowers
Perianth Perianth segments in the form of numerous slender threads or elongated spoon-like
scales.
Androecium Male flowers with 2–5 stamens; variously monadelphous; filaments free or connate;
connate filaments bear long silky hairs; anthers basifixed, dithecous; connective projecting beyond
the anthers.
Gynoecium Absent.
Female Flowers
Perianth Same as that of male flowers.
Androecium Absent.
Gynoecium Monocarpellary, superior, unilocular, 1 seeded; present on a stipe bearing slender hairs
and containing 1 pendant ovule; style 1, long; stigma 1, linear to spathulate or rhomboidal.
Fruit An achene bearing the persistent style and usually surrounded by persistent scales.
Seeds With mealy endosperm; embryo long or straight and narrow.
Floral Formulae
(a) Male Flower: Br, ≈, , Pμ, A(2 – 5), G0.
(b) Female Flower: Br, ≈, , Pμ, A0, G1.

• Dried leaves of Typha are used as weaving materials in making mats, chair bottoms, baskets,
etc.
• Dried, dense cylindrical spikes of inflorescences are used in floral arrangements.
• Occasionally, some species of Typha are grown as ornamentals in ponds and river
margins.
• Leaf bases and rhizomes are eaten by mammals including humans in some parts of India.
Bur, a yellow cake prepared from the flowers of T. angustata, is eaten by natives.
• Rootstocks of Typha angustata are astringent and diuretic.

Typhaceae has been included under series Nudiflorae by Bentham and Hooker along with 4 more fam-
ilies, viz. Araceae, Pandanaceae, Lemnaceae and Cyclanthaceae. Typhaceae has however been treated
by Engler and Prantl under order Pandanales along with one more family, Pandanaceae. Benson also
treated Typhaceae under Pandanales as treated already by Engler and Prantl. Hutchinson, however,
discussed Typhaceae under order Typhales along with one more family, namely Sparganiaceae.
Modern taxonomists like Hutchison (1964), Takhtajan (1980) and Cronquist (1981) consider
Typhaceae as an advanced family of monocotyledons because of several characters, including (i) exs-
tipulate leaves with sheathing bases, (ii) flowers arranged in dense spikes, (iii) unisexual flowers,
(iv) reduced perianth in the form of hair-like outgrowth’s, and (v) monocarpellary gynoecium.
22.14 ARACEAE ARUM FAMILY

Monocotyledons, Nudiflorae, Araceae.

Rhizomatous or tuberous herbs, found in both aquatic and terrestrial habitats; inflorescence spadix
subtended or enveloped by a single spathe; flowers small and often bad smelling; fruit a berry.

Acorus, Alocasia, Amorphophallus, Colocasia, Pistia, Pothos.

A family of 115 genera and over 2000 species, Araceae are distributed mostly in tropical and some
also in temperate regions of the world. About 25 genera and over 140 species have been reported
from India, chiefly from western and southern parts. Some larger genera along with the number
of their worldover reported species are Anthurium (500), Philodendron (275), Arisaema (150),
Amorphophallus (100), Rhaphidophora (100), Pothos (75), Alocasia (70), Monstera (50), Arum
(15), Colocasia (8), Acorus (2) and Pistia (1). Colocasia (Colocasia antiquorum) and ‘money plant’
(Pothos aureus) are two famous plants of Araceae.

General Habit Highly variable; usually perennial, rhizomatous or tuberous herbs (Arum), climbers
(Pothos), or tree-like (Dracontium, Philodendron); rarely free-floating aquatic herbs (Pistia); some
are epiphytes (Anthurium), or occur in marshy conditions (Lasia); often raphides or calcium oxalate
crystals present in the sap of the plants.
Root Fibrous, adventitious, and usually of two types i.e. (i) climbing or clasping roots, which cling
to the support, and (ii) absorbing roots which enter into the soil and absorb nutrients; velamen
develops in the aerial roots of epiphytic species.
Stem Subterranean or underground in the form of tubers (Arum), corms (Amorphophallus), rhizome
(Acorus) or root stocks (Colocasia); or aerial (Pothos) showing monopodial branching or sympodial;
accessory buds often develop in leaf-axils.
Leaf Variable in shape and types; single (Arisaema) or a few; basal (Arisaema), or cauline and
alternate (Pothos); often hastate (Typhonium), sagittate or round; pinnately or palmately divided; peti-
olate (Philodendron) or sessile (Pistia); venation is usually parallel but rarely reticulate (Arum).
508 Plant Taxonomy
Inflorescence Flowers usually grouped together to form a more

or less cylindrical spadix subtended by a large coloured spathe;
the upper portion of spadix is usually naked and the lower por- spathe
tion bears whorls of small unisexual flowers, the male flowers
above and the female below (Fig. 22.19); usually some sterile
male and female flowers are present close to fertile whorls of
male and female flowers; male and female flowers develop on upper portion
different spadix in some genera, e.g. Arisaema. of spadix
Flower Ebracteate, ebracteolate, usually unisexual but some-

times bisexual (Acorus, Pothos), actinomorphic, hypogynous or
epigynous, dimerous or trimerous, small, often bad-smelling.
Perianth Usually absent in unisexual flowers (Arum, Pistia)
and present in bisexual flowers (Acorus, Fig. 22.20); if pres- male flowers
(sterile)
ent, the perianth lobes are small, scale-like, 4 to 6 in number,
present generally in 2 whorls of 2 + 2 or 3 + 3, free or rarely
connate (Spathiphyllum). male flowers
Androecium Usually 6 or lesser stamens present in one or (fertile)
2 whorls, located opposite the perianth lobes; free or united
female flowers
into a synandrium (Colocasia, Spathicarpa); anthers dithecous, (sterile)
introrse, dehiscing by slits or pores; female flowers usually
bear staminodes. female flowers
Gynoecium Varied in structure but often reduced to a single (fertile)
carpel; sometimes 2 to 9 united carpels; superior or rarely
inferior and embedded in the spadix; one to many locules and
one to numerous ovules in each locule; placentation is basal
(Arisaema, Typhonium), axile (Pothos) or parietal (Arum); style Fig. 22.19 L.S. inflorescence
various, sometimes greatly reduced; stigma variable. of Arum maculatum.
Fruit and Seeds Fruit usually a berry, clustered closely on the
spadix and appearing as a multiple fruit. Seeds with or without
endosperm.
Pollination Flowers are protogynous and insect-pollinated.
Insects are attracted because of the coloured spathe and charac-
teristic bad smell emitted by inflorescences. Insects come into
contact with the fertile female flowers and transfer to them any
pollen they may be carrying from the male flowers of another
plant. Some species of Arisaema show self-pollination.
(a) Male Flowers: Ebr, ≈, , P0, A3+3 or 2+2, G0.
(b) Female Flowers: Ebr, ≈, , P0, A0, G1 or (2–9).
(c) Bisexual Flowers: Ebr, ≈, , P3+3 or 2+2, A3+3 or 2+2, Fig. 22.20 Floral diagram
G1 or (2–9). of Acorus.

• Widely used vegetables are obtained from (i) the stem and root-stocks of Alocasia indica
(‘Mankanda’), (ii) tuberous corms and leaves of Amorphophallus campanulatus (‘Zimikand’
or ‘elephant foot’), (iii) tubers of Colocasia esculenta syn C. antiquorum (‘Arvi’, ‘Ghuiyan’,
‘Kachalu’ or Colocasia), and (iv) corms and leaves of Remusatia vivipara and Xanthosoma
niger.
• Industrial alcohol and starchy baby foods are also prepared from the tubers of Colocasia
esculenta.
• Certain plants are of medicinal value. These include (i) Acorus calamus, of which the rhi-
zomes are beneficial in chronic diarrhoea and dyspepsia, (ii) Alocasia macrorrhiza, of which
the stem juice is used to relieve pain from scorpion bite, (iii) Amorphophallus campanulatus,
of which the corms are used in treating piles and dysentery, and (iv) Scindapsus officinalis,
of which the dried fruits have anthelmintic properties.
• Plants of ornamental value, grown commonly in gardens for their often variegated and strik-
ingly handsome leaves are Acorus calamus var. variegatus, Alocasia indica var. metallica,
Caladium bicolor, C. picturatum, Colocasia antiquorum, Dieffenbachia picta, Monstera
deliciosa, Pothos aureus (Money plant), Scindapsus officinalis and S. pictus.
• Ornamental plants cultivated mainly for their flowers and spathes include several species
of Anthurium, Arisaema (jack-in-the-pulpit or Indian turnip), Dracunculus (dragon arum),
Lysichiton (western skunk cabbage), Sauromatum (Voondoo lily) and Spathiphyllum.
• Pistia stratioites is cultivated for being used in aquaria.

Placed under series Nudiflorae by Bentham and Hooker, family Araceae was included in order
Spathiflorae in Engler’s system, and was placed under order Arales by most of the other taxonomists
including Hutchinson, Takhtajan, Cronquist, Thorne, etc. In Engler’s system, the family was divided
into eight subfamilies viz. Pothoideae, Monsteroideae, Calloideae, Lasioideae, Philodendroideae,
Colocasioideae, Aroideae and Pistioideae.
Takhtajan (1969) includes only two families in Arales viz. Araceae and Lemnaceae. He opined
that Arales “most likely has a common origin with the Arecales and Cyclanthales from intermediate
ancestors of the Liliales.”
In affinities, Araceae appears to be closely related to Palmae, because both families have same
type of inflorescence, and small, hypogynous, actinomorphic flowers.
22.15 ALISMATACEAE OR ALISMACEAE WATER PLAINTAIN FAMILY

Monocotyledons, Apocarpae, Alismataceae.
510 Plant Taxonomy

Aquatic or marshy herbs with perennating rhizomes; bisexual flowers; perianth biseriate with 3 free
sepals and 3 free petals; many free carpels.

Alisma, Limnophyton, Sagittaria.

Represented by about 14 genera and over 90 species, Alismataceae are cosmopolitan in distribution.
About 7 genera and over 10 species have been reported from India. The chief genera along with
the number of their reported species and common names of some are Echinodorus (25, burhead),
Sagittaria (20, arrowhead), Alisma (10, water plaintain), and Damasonium (5).

General Habit Annual or perennial, aquatic or marshy herbs with a stout perennating rhizome; roots
are fibrous, and stem is thin, erect or with floating leaves; in Sagittaria the runners terminate into
tubers; usually the latex is present in stem and leaves.
Leaf Radical or clustered at nodes, long-petioled, leaf base sheathing; erect, floating or submerged;
blades ovate or linear to lanceolate with bases sometimes sagittate or hastate; parallel venation;
usually small scales develop in axils.
Inflorescence Usually much branched or whorled; primary branches often in racemose racemes or
panicles, the secondary branches often cymose.
Flower Bracteate (Butomus) or ebracteate, pedicellate; bisexual (Alisma, Fig. 22.21; Butomus) or
unisexual (Sagittaria); actinomorphic, trimerous, hypogynous with a flat, convex or dome-shaped
receptacle.
Perianth Biseriate, consisting of 6 perianth lobes arranged usually in 2 whorls of 3 lobes; the outer
whorl, representing the calyx, consists of 3 free, green sepals which are twisted (Alisma, Fig. 22.22),
valvate (Sagittaria, Fig. 22.23) or imbricate (Butomus); the inner whorl, representing the corolla,
consists of 3 free, coloured petals which are also twisted, valvate or imbricate.
Androecium Stamens are usually 6 (Alisma) but variable between 3 (Wisneria) to numerous
(Sagittaria) in different genera; 9 stamens in Butomus are arranged in an outer whorl of 6 and
inner whorl of 3 stamens; filaments free and anthers are dithecous, basifixed and extrorse in most
of the genera.
Gynoecium Carpels are usually 6 and apocarpous but variable from 3 (Wisneria) to numerous
(Alisma) free carpels; ovary of each carpel is superior, unilocular, with a single ovule, basal placen-
tation; rarely the ovules are 2 or more in each locule; style one, persistent, bear stigmatic papillae
(Alisma, Fig. 22.21).
Fruit and Seeds Fruit a group of achenes arranged spirally or in ring. Seeds with a horse-shoe
shaped or curved embryo and without endosperm.
stigma style stamen

stamen
petal
carpels
petal sepal
carpel
A Flower
sepal
ovary
ovary
L.S. Flower
locule
ovule
inflorescence
Stamen
stigmatic papillae
stigma
L.S. Carpel
style
Style
(upper portion)
Flowering Plant
Fig. 22.21 Alisma plantago-aquatica L.

512 Plant Taxonomy
Fig. 22.22 Floral diagram of Alisma.
A B
Fig. 22.23 Floral diagrams of Sagittaria guayanensis—A: Male flower; B: Female flower.
Pollination and Dispersal Flowers are entomophilous, and get pollinated by flies, short-tongued bees
and other similar insects which visit them in search of nectar. Dispersal of seeds takes place mainly
by water and partly by air.
Floral Formula (Bisexual Flower) Br or Ebr, ≈, , K3, C3, A6, G6 or 3–μ.

Economically, the family is not of any special significance. Sagittaria rhizomes are sometimes
grown for their food value and used as food under the name “swamp potato”. Tubers of Alisma
plantago-aquatica and corms of Sagittaria sagittifolia are edible. Some species of Sagittaria and
Alisma are used as aquarium plants. Some species of Alisma, Echinodorus and Sagittaria are
planted for decoration besides ponds and streams.

Family Alismataceae was included under series Apocarpae by Bentham and Hooker, order Helobiae
by Engler, order Alismales by Takhtajan, and under order Alismatales by Cronquist, and also by
Thorne. Usually, the family is divided into two subfamilies viz. Alismatoideae (ovules solitary in
each carpel) and Butomoideae (ovules many in each carpel).
Alismataceae is closely related to Ranunculaceae of dicotyledons in possessing hypogynous flow-
ers, biseriate perianth differentiated into calyx and corolla, free sepals and petals, and often indefi-
nite number of free stamens and carpels. Because of these similarities, certain theories of origin of
monocots from dicots suggest Alismataceae as the connecting link between the two.
Brown (1875) believed that most primitive flowers were those of Magnoliaceae and Alismaceae.
However, Takhtajan (1969) stated that Alismales “exhibits definite links with the Nymphaeales
among the dicots”, and the flowers of both are primitive. However, the seeds are endospermic in
Nymphaeales and without endosperm in Alismales. Because of several close similarities between
Alismatales and dicotyledons, Cronquist (1981) discussed order Alismatales in the very beginning
of Liliopsida (Monocots).
22.16 CYPERACEAE SEDGE FAMILY

Monocotyledons, Glumaceae, Cyperaceae.

Herbs with triangular stem; leaves 3-ranked, ligule absent; flowers in spikelets; bracts glumaceous;
lodicules absent; fruit an achene or nut.

Bulbostylis, Carex, Cyperus, Eleocharis, Fimbristylis, Scirpus.

A family of approximately 90 genera and over 4000 species, showing worldwide distribution in
damp and wet habitats. About 23 genera and over 400 species have been reported from India. Larger
genera of the family, along with the number of their approximately reported species from the world
514 Plant Taxonomy
and the common names of some of them are Carex (1100, Carex or sedge), Cyperus (700, sedge),
Fimbristylis (300), Scirpus (250, bulrush), Rhynchospora (225, beak rush), Eleocharis (200, spike
rush), Scleria (200, nut rush), Schoenus (100) and Uncinia (35). Cyperus papyrus, used for paper-
making by Egyptians even as early as 2400 B.C., and C. rotundus, a serious weed of lawns and
fields, are two important plants of the family.

General Habit Annual or perennial, grass-like herbs found in damp or wet places; persisting usually
by a creeping, underground, sympodial rhizome.
Roots Fibrous, developing from the underground rhizome or tuberous stem.
Stem Often called ‘culm’, the aerial stem is often triangular, solid or rarely hollow, leafless and
generally unbranched below the inflorescence.
Leaf Grass-like, often develop in a crowded tuft on the lower basal part of the culm, three-ranked (in
1/3 phyllotaxy), with a closed sheath; leaf blades long, narrow, with a thick midrib; ligule absent.
Inflorescence Basic unit of the inflorescence is a spikelet (Fig. 22.24); the spikelets are usually 1 to
many-flowered and variously arranged in racemes, panicles, spikes, or umbels, and usually subtended
by well-developed leaf-like bracts; large bracts are distichously or spirally arranged.
Flower Develop singly in the axil of glumes; unisexual (Carex, Fig. 22.25) or bisexual (Cyperus,
Fig. 22.26); trimerous, hypogynous; small or inconspicuous; unisexual species may be monoecious
or dioecious; the female flower in some genera (Carex) is enclosed by a modified glume called
utricle or perigynium (Fig. 22.25B).
Perianth Absent (Cyperus) or represented by usually 6 hypogynous bristles (Scirpus), scales, or
hairs (Fig. 22.27A).
Androecium Stamens are generally 3 (Cyperus rotundus), less commonly 2 (C. alulatus, C. iria)
or 1 (C. squarrosus), and rarely 6 (Gahnia); filaments free; anthers basifixed, dithecous, introrse;
Gynoecium Bi- to tricarpellary, syncarpous; ovary superior, unilocular, with 1 anatropous ovule;
basal placentation; ovary in Carex is subtended and enclosed in a perigynium (Fig. 22.25B); styles
2 to 3 or deeply divided into as many branches as the number of carpels.
Fruit and Seeds Fruit a triangular or biconvex achene or nutlet. Seed one per fruit, with small
embryo, embedded in fleshy or mealy endosperm; seed coat free from the pericarp.
Pollination Flowers are usually wind-pollinated, although pollen-eating beetles may also transport
pollen grains in some species of Carex. Several Carex species are protandrous.
(a) Male Flowers: ≈, , P0 or bristles, A1–3, G0.
(b) Female Flowers: ≈, , P0 or bristles, A0, G (2–3).
(c) Bisexual Flowers: ≈, , P0 or bristles, A1–3, G (2–3).
LB
C D
SP
SP
E F
Fig. 22.24 Inflorescence of a few species of Cyperus. A: C. compressus; B: C. globosus All. syn. Pycreus
globosus Reich; C: C. metzii (Hochst) Mattf. syn. Kyllinga squamulata Than; D: C. triceps (Rottb.) Endl. syn.
Kyllinga triceps Rottb; E: C. triceps showing five spikes; F: C. paniceus (Rottb.) Boeck. var. roxburghianus
syn. Mariscus paniceus Vahl. var. roxburghianus Clarke (C, Culm; LB, Leafy bract; S, Spikelet; SP, Spike)
(Photographs from author’s Ph. D. Thesis).
516 Plant Taxonomy
A B
Fig. 22.25 Floral diagrams of Carex cruciata—A: Male flower; B: Female flower (Note the perigynium
in ‘B’).
Fig. 22.26 Floral diagram of Cyperus rotundus.

Sedges are not of much significant importance. However, some are grown as ornamentals,
few are used in medicines, and some are even edible. Several of them are noxious weeds
(Cyperus rotundus) of lawns and cultivated fields. Some aspects of their economic importance
are undermentioned:
• Starchy tubers of some members are edible, e.g. Cyperus esculentus (‘Chufas’ or Tiger-nut),
C. bulbosus, and Eleocharis dulcis (Chinese water-chestnut).
stamen
perianth
of bristles
stamen
pistil pistil
bract
A B
Fig. 22.27 Vertical view of flowers of Scirpus (A) and Cyperus (B).
• Tubers of Cyperus rotundus are used in stomach disorders, while that of C. articulatus are
used as tonic, and that of C. stoloniferous are considered as stimulant for the heart. Tubers
of C. scariosus are used in treating stone trouble of kidney and urinary bladder.
• Chufa-oil, obtained from the tubers of Cyperus esculentus, is used in soap industry.
• Tubers of Cyperus rotundus and C. scariosus are used in the preparation of fragrant sticks
called “Agarbattis” in India.
• Cyperus papyrus (Papyrus plant), formerly used in paper-making, is nowadays grown as an
ornamental plant. Other plants of ornamental value are C. flabelliformis syn. C. alternifolius
(umbrella plant), C. fertilis and C. gracilis.
• Culms of Cladium mariscus, Cyperus corymbosus, C. elatus, C. exaltatus, C. iria, etc. are
used for making mats, baskets, etc.
• Several species of Cyperus, Carex, Scirpus, Rhynchospora, etc. are used as fodder, though
of inferior quality.

Taxonomists generally treat Cyperaceae to be closely allied to Gramineae, and that is why Bentham
and Hooker treated both of them under the same series Glumaceae, and the two families have been
included under the same order Glumiflorae by Engler as well as Rendle. Cronquist (1981) also
included both of them under order Cyperales, and the two families have been included under one
and the same order Commelinales by Thorne (1983). Hutchinson (1959), however, put them under 2
monotypic orders, i.e. Cyperaceae under Cyperales and Gramineae under Graminales, and according
to him both the families have been derived from Juncales but from different stocks on parallel lines.
Takhtajan (1969) opined that Cyperaceae, the only family of order Cyperales, “evidently derived
directly from the most primitive Juncaceae”.
518 Plant Taxonomy
Hickey and King (1988) divided Cyperaceae into 3 subfamilies viz. (1) Cyperoideae (flowers bisex-
ual, spikelets many-flowered, rarely flowers unisexual; e.g. Cyperus, Scirpus), (ii) Rhynchosporoideae
(flowers unisexual or bisexual, with or without perianth; in few-flowered, spike-like cymes grouped
into heads or spikes; e.g. Rhynchospora), (iii) Caricoideae (unisexual flowers, without perianth;
usually in many-flowered spikes; perigynium present; e.g. Carex, Uncinia).
Radford (1986) mentioned 8 tribes under Cyperaceae, viz. 1. Cypereae (Cyperus), 2. Dulichieae
(Dulichium), 3. Rhynchosporeae (Cladium), 4. Scirpeae (Scirpus) 5. Hypolytreae (Mapania), 6.
Sclerieae (Scleria), 7. Cryptangieae (Cryptangium), and 8. Cariceae (Carex).
22.17 GRAMINEAE OR POACEAE GRASS FAMILY

Monocotyledons, Glumaceae, Gramineae.

Herbs with round stem (culm) having usually hollow internodes; leaves mostly flat, 2-ranked and
usually with open leaf sheath; ligule usually present; bracts of glumes, lemma and palea; fruit
caryopsis.

Avena, Bambusa, Cynodon, Hordeum, Oryza, Poa, Saccharum, Secale, Sorghum, Triticum, Zea.

Gramineae, the largest and most important family of angiosperms from the economic point of view,
is represented by about 620 genera and over 10,000 species, and distributed widely in almost all
regions of the world and in almost every type of habitat. About 240 genera and over 1200 species
have been reported from India. Some of the largely represented genera along with the total number
of their species and common names of some of them are Panicum (500, panic grass), Digitaria
(380, crab grass), Aristida (330, wire grass), Poa (300, blue grass), Eragrostis (300, love grass),
Stipa (300, needle grass), Paspalum (250, Bahia grass), Agrostis (150–200, bent grass), Arundinaria
(150, giant cane), Pennisetum (150, Bajra), Setaria (140, Italian millet), Agropyron (100–150, wheat
grass), Muhlenbergia (100, bull grass), Avena (70, oats), Sorghum (60, sorghum), Oryza (25, rice),
Triticum (20, wheat), Hordeum (20, barley), Saccharum (12, sugarcane), Cynodon (10, Doob grass),
Zea (1, maize).

General Habit Usually annual or perennial herbs; rarely woody shrubs or tree-like, reaching up to
a height of 30 metre (Bambusa) or even more.
Roots Fibrous and adventitious; primary roots are often short-lived; some adventitious roots (stilt
roots) in Zea mays develop from the basal nodes of stem for providing extra support.
Stem The stems (‘culms’) are erect, ascending, prostrate or creeping; simple or commonly branched
at the base; usually round, jointed, and bear hollow internodes, and solid and swollen nodes; termi-
nates usually by inflorescence; many species have a perennial, creeping rhizome or stolon formed
by the lower internodes of stem.
Leaf Simple, alternate, in two rows on opposite sides of the stem (2-ranked), originating at the
nodes, often crowded at the base, and consisting of sheath, blade and ligule; sheath with margins
free and overlapping or encircling the culm to form a tube; blade or lamina usually long, narrow,
flat, linear to lanceolate, rarely with a constricted petiole-like base (Bambusa); veins parallel; ligule,
present at the junction of blade and sheath on the adaxial surface, is membranous, or reduced to a
ring of hairs, or even absent (Echinochloa).
Inflorescence Basic unit of inflorescence is a spikelet
(Fig. 22.28); each spikelet consists of one or more flowers (or
florets) and their subtending bracts arranged on an inflores-
cence axis called rachilla; spikelets are either sessile or with
pedicel, and arranged in spikes, racemes or panicles forming
compound inflorescences; at the base of each spikelet are
generally present two sterile glumes, of which the lower one
is called first glume and the upper one is called second glume;
in some genera, either the first glume or both the first and
second glumes are absent; up to 6 sterile glumes are present
in some genera; the flowers are arranged on the rachilla just rachilla
above the second glume. stamen
stigma
Flower or Floret Flowers, also called florets (Fig. 22.29), are
small, inconspicuous, bisexual or unisexual (Zea, Fig. 22.30), ovary
zygomorphic, hypogynous; develop on the rachilla above the
glumes; and each floret is subtended by two bracts called
lemma and palea; lemma, also called fertile or flowering palea
glume, is greenish, keeled, nerved or awned, and often resem- lemma
bles with sterile glumes; palea, present in between rachilla and
lemma, is thin, membranous, 2-nerved or 2-keeled, and often lodicule
partially enclosed by the lemma.
Palea, morphologically representing a bracteole, bears a
flower proper in its axil.
Perianth Absent (Dendrocalamus) or reduced to usually 2 or second first
glume glume
rarely 3 minute scales called lodicules; lodicules are fleshy or
hyaline and present antero-laterally, if 2 in number.
Androecium Stamens 1 to 6 or rarely more, but usually 3 in Fig. 22.28 Diagrammatic
each floret (Fig. 22.31); the odd stamen is always anterior; representation of a spikelet of
only 1 anterior stamen is present in Uniola; filaments free; Triticum aestivum.
anthers dithecous, basifixed or versatile, introrse, and usually
opening by a longitudinal slit.
520 Plant Taxonomy
awn
palea
stigma
rachilla
ovary
lodicule
lemma
with awn
filament
glume anther
Fig. 22.29 Parts of a bisexual flower of Gramineae.
Gynoecium Bi- or tricarpellary, syncarpous; ovary superior, unilocular, containing one anatropous
ovule, basal placentation; styles 1 (Nardus) to 3 (Bambusa), commonly 2; stigmas typically 2, often
plumose or feather-like.
Fruit and Seeds Fruit usually a caryopsis or rarely a nut, utricle (Sporobolus) or berry (some
bamboos). Seed 1 per fruit, with abundant and starchy endosperm; embryo variable in size.
Pollination Plants, such as Triticum, are self-pollinated whereas majority of grasses are wind-
pollinated. Flowers are protogynous. Characters, such as small and inconspicuous flowers, and
feather-like stigma, are the adaptations for anemophily. Flowers produce loose and powdery pollen
grains in large quantity.
General Floral Formula ≈ or , or or , P2 or 3 or absent, A3 or 1–6, G (2–3).

Gramineae are of greater importance than any other family of flowering plants from the economic
point of view. Plants of this family provide food for man (wheat, oat, barley, sugarcane, etc.),
fodder for our domestic animals, grasses and ornamental plants for our lawns and gardens, and
several other products including starch, ethyl alcohol, paper, fibres, edible oils, adhesives, thatch-
ing and building materials, wildlife food, etc.
• Cereals1
(i) Avena sativa (Oats, Jai): This important cereal and fodder crop is highly nutritious.
(ii) Hordeum vulgare (Barley or Jaun): Used as a feed grain and also in manufacturing
several alcoholic beverages including beer, barley water, etc.
1
styles
palea
lemma
glume
stamen
rachilla
B
C
pistillate
florets glume
lemma style
axis
ovary
floral axis hairs

D
embryo
remains of
style
A
endosperm
E
pericarp
Fig. 22.30 Zea mays: A: L.S. female inflorescence; B: A spikelet of 2 male florets, of which one is at
anthesis; C: L.S. of two male florets (only nearest stamens shown); D: L.S. female floret; E: L.S. caryopsis
surrounded by other floral parts.
(iii) Oryza sativa (Rice, Chawal or Paddy): Used as a common staple food by millions
of people throughout the world, rice is also used as a livestock and for manufacturing
paper, alcohol, starch etc. Its husk is used as fuel and its bran is the source of fatty
oil.
(iv) Secale cereale (Rye): Besides its limited use as a cereal crop, rye is used for making
alcoholic beverages, pickles, sambar, etc.
(v) Triticum aestivum syn. T. vulgare (Wheat, Gehun): Wheat is used universally as a
staple food for man. Its straw is used as stockfeed for cattles, for packing goods, and for
522 Plant Taxonomy
palea
lodicule
lemma
Fig. 22.31 Floral diagram of Triticum aestivum.
manufacturing paper. Grains are also used for manufacturing starch, industrial alcohol,
etc.
(vi) Zea mays (Maize, Makka): Used as a common food by man, maize is also used as
a livestock feed for cattles and also for manufacturing several industrial products such
as explosives, adhesives, soaps, starch, plastics, asbestos, linoleum, etc. Paper is manu-
factured from its leaf stalks, and the corn oil is used in preparing paints, varnishes,
etc. Several alcoholic beverages are also manufactured from maize.
• Millets: Millets1, used as coarse grains by man and common fodder for animals, belong
to Gramineae. Some of the important millets include Eleusine coracana (Ragi or finger
millet), Panicum miliaceum (sama or common millet), Pennisetum typhoides (Bajra or pearl
millet), Setaria italica (Italian millet), and Sorghum vulgare (Jowar or sorghum). Jowar and
Bajra are chiefly used for poultry and cattles, and also for preparing alcoholic beverages
and diastase.
• Sugar: It is obtained from the stem juice of Saccharum officinarum (sugarcane or Ganna).
Baggase is used for manufacturing paper while the molasses is utilized for manufacturing
industrial alcohol and alcoholic beverages such as rum.
• Fodder: The whole aerial green part of several members of Gramineae is cut and used
as green fodder. Several species of some commonly used genera for this purpose belong to
Avena, Eleusine, Hordeum, Pennisetum, Phalaris, Setaria and Sorghum.
1
Almost all wild grasses are browsed by the cattles, of which some commonly browsed
species are Andropogon pertusus, Apluda mutica, Cenchrus ciliaris, Cynodon dactylon,
Dichanthium annulatum, Iseilema laxum and Panicum maximum.
Straw of various species of Oryza and Triticum is also used as a common fodder for
cattles.
Some other fodder grasses are Dactylis glomerata, Heteropogon contortus, Lolium multi-
florum, Paspalum scrobiculatum, Poa annua, Sporobolus diander and Stipa orientalis.
• Lawn Grasses: Many grasses are used in lawns and other turfed areas, e.g. Agrostis,
Cynosurus, Festuca, Lolium, and Poa.
• Paper Manufacturing: Pulp, prepared from the straw of several species of this family, is
used in manufacturing paper of almost all coarse and fine qualities, strawboard, artificial
rayon, etc. Some commonly used genera for the purpose are Bambusa, Cephalostachyum,
Dendrocalamus, Erianthus (E. arundinaceus, E. munja, E. ravennae), Gigantochloa,
Melocanna, Narenga, Ochlandra, Oryza, Saccharum (S. benghalense, S. officinarum, S.
procerum, S. spontaneum), Triticum aestivum, Vetiveria zizanioides and Zea mays.
• Bamboos: ‘Bamboos’, a common term used for some tall tree-like woody grasses, are
universally used for papermaking. They are also used for house building, as substitute of
timber, and also for preparing several other articles such as walking sticks, beds, tent poles,
umbrella handles, kites, musical instruments, furnitures, toys, mats, etc. Some bamboos
are harvested for cattle fodder, and several others are grown for checking soil erosion,
and also for ornamental purposes. Some common bamboos are Arundinaria, Bambusa,
Cephalostachyum, Dendrocalamus, Melocanna, Ochlandra and Phyllostachys.
• Khus-Khus: Roots of Vetiveria zizanioides are used for making curtains and screens
(Khus-Khus), which, when wetted, bring about cooling and scenting of rooms.
• Essential Oils: Volatile oil, obtained from the leaves of several grasses, is used in per-
fumery, medicines, mosquito-repellent creams; soaps, etc. Some grasses utilised for the
purpose are Cymbopogon caesius (ginger grass oil), C. citratus (lemon grass oil), C. mar-
tinii (palmarosa oil), C. nardus (Citronella oil) and Vetiveria zizanioides (khus oil or oil of
vetiver).
• Ornamental Plants: Grasses, grown for ornamental purposes in the gardens, or dried and
used for floral decoration include Axonopus affinis (carpet grass), Briza maxima (quaking
grass), Coix lacryma-jobi (job’s tears), Cortaderia selloana (Pampas Grass), Cynodon dac-
tylon (Doob grass or Bahama grass), Lagurus ovatus (Hare’s-tail grass), Lamarckia aurea
(golden top grass), Phalaris arundinacea (ribbon grass), P. canariensis (canary grass),
Setaria italica (Foxtail millet) and Stipa pinnata (feather grass).
• Brooms: Panicles of Thysanolaena maxima (Broom grass) are used as soft brooms.
• Ropes, Mats, Baskets: Ropes are made from the fibres obtained from the leaves of
Erianthus munja syn. Saccharum munja (Munj) and Saccharum spontaneum (Kaans) while
mats, baskets, musical instruments, etc. are prepared from the stems of Arundo donax
(Nal or reed), Arundinaria falcata, A. racemosa and Phragmites karka (Bansi or common
reed).
524 Plant Taxonomy

Family Gramineae or Poaceae has been variously placed under series Glumaceae by Bentham
and Hooker (1862–1883), order Glumiflorae by Engler and Prantl (1931), order Graminales by
Hutchinson (1959), order Poales by Takhtajan (1969), order Cyperales by Cronquist (1981), and order
Commelinales by Thorne (1983).
Family Gramineae is usually divided into 6 subfamilies, divisible further into 25 tribes as
under:
Subfamily 1. Festucoideae: Tribe 1. Festuceae (Poa); Tribe 2. Aveneae (Avena); Tribe 3.
Triticeae (Triticum); Tribe 4. Meliceae (Melica); Tribe 5. Stipeae (Stipa); Tribe 6. Brachyelytreae
(Brachyelytrum); Tribe 7. Diarrheneae (Diarrhena); Tribe 8. Nardeae (Nardus); Tribe 9. Monermeae
(Monerma).
Subfamily 2. Panicoideae: Tribe 10. Paniceae (Pennisetum); Tribe 11. Andropogoneae
(Saccharum).
Subfamily 3. Eragrostoideae: Tribe 12. Eragrosteae (Eragrostis); Tribe 13. Chlorideae (Cynodon);
Tribe 14. Zoysieae (Zoysia); Tribe 15. Aleuropodeae (Allolepis); Tribe 16. Unioleae (Uniola); Tribe
17. Pappophoreae (Cottea); Tribe 18. Orcuttieae (Orcuttia); Tribe 19. Aristideae (Aristida).
Subfamily 4. Bambusoideae: Tribe 20. Bambuseae (Bambusa); Tribe 21. Phareae (Pharus).
Subfamily 5. Oryzoideae: Tribe 22. Oryzeae (Oryza).
Subfamily 6. Arundinoideae: Tribe 23. Arundineae (Arundo); Tribe 24. Danthonieae (Danthonia);
Tribe 25. Centotheceae (Chasmanthium).
Superficially, Gramineae resemble Cyperaceae. In both these families, the flowers are small,
inconspicuous, subtended by bracts, and present in the form of spikelets. However, Gramineae differ
from Cyperaceae by their usually terete, hollow, jointed stems, distichous leaves, and structure of
spikelet. Hutchinson opined that the Gramineae have been derived from Restionaceae of Juncales
which again have been derived from Liliaceous stock. Some are also of the opinion that Gramineae
may be distantly related to the Commelinaceae.

1. Following genera belong to which family of monocotyledons?
1. Zeuxine 2. Vanda 3. Gladiolus 4. Crocus
5. Agave 6. Ananas 7. Canna 8. Ravenala
9. Curcuma 10. Elettaria 11. Aloe 12. Asparagus
13. Ruscus 14. Tradescantia 15. Phoenix 16. Colocasia
17. Pothos
2. Describe floral structure of family Arecaceae (Palmae) in semitechnical language.
3. Describe the basic structure of the spikelet of Cyperaceae.
4. Discuss the inflorescence and floret details of Poaceae.
5. “Poaceae (Gramineae) are of greater economic importance than any other family of flower-
ing plants”. Comment.
Suggested Reading
Bentham, G. and J.D. Hooker, 1862–1883, Genera Plantarum, 3 Vols., Reeve & Co., London.
Cronquist, A., 1981, An Integrated System of Classification of Flowering Plants, Columbia Univ. Press,
New York.
Engler, A.W. and K. Prantl, 1887–1915, Die Naturlichen Pflanzenfamilien, 23 Vols., Leipzig.
Heywood, V.H., 1978, Flowering Plants of the World, Mayflower Books, New York.
Hickey, M. and C. King, 1988, 100 Families of Flowering Plants, (2nd ed.), Cambridge University Press,
Cambridge.
Holmes, S., 1983, Outline of Plant Classification, Longman, London.
Hutchinson, J., 1969, Evolution and Phylogeny of Flowering Plants, Academic Press, London.
________ 1973, The Families of Flowering Plants, (3rd ed.), Clarendon Press, Oxford.
Jones, S.B. Jr. and A.E. Luchsinger, 1987, Plant Systematics, (2nd ed.), McGraw-Hill Book Co., New
York.
Lawrence, G.H.M., 1951, Taxonomy of Vascular Plants, Macmillan Co., New York.
Radford, A.E., 1986, Fundamentals of Plant Systematics, Harper & Row Publishers, Inc., New York.
Rendle, A.B., 1925, The Classification of Flowering Plants Vol. II (Dicotyledons), Cambridge Univ. Press,
London.
Takhtajan, A., 1969, Flowering Plants: Origin and Disperal, Oliver & Boyd Ltd., Edinburgh.
________ 1980, Outline of the classification of flowering plants (Magnoliophyta), Bot. Rev. 46: 225–359.
Thorne, R.F., 1983, Proposed new realignments in the angiosperms, Nordic J. Bot. 3: 85–117.
Willis, J.C., 1973, A Dictionary of the Flowering Plants and Ferns, (8th ed., revised by H.K. Airy Shaw),
Cambridge Univ. Press, London.
APPENDIX 1
EXAMINATION TOOLS COMPARATIVE TABLES OF SELECTED FAMILIES
Table A.1 Magnoliaceae and Annonaceae
Magnoliaceae Annonaceae
1. Leaves usually stipulate. 1. Leaves usually exstipulate.

2. Usually the sepals and petals are undifferentiated. 2. Sepals and petals are clearly differentiated.
3. Anthers are usually introrse. 3. Anthers are usually extrorse.
4. Style and stigma absent in some species. 4. Style and stigma usually present.
5. Sarcotesta present in the seeds. 5. Sarcotesta absent.
6. Wood is not usually aromatic. 6. Wood usually is aromatic.
Table A.2 Ranunculaceae and Nymphaeaceae
Ranunculaceae Nymphaeaceae
1. Usually terrestrial, and only rarely aquatic. 1. Mostly aquatic.

2. Plants do not possess latex. 2. Plants usually possess colourless or coloured latex.
3. Stem is usually aerial, erect or climbing; only 3. Stem is mostly a rhizome, usually burried
few possess rhizome. in mud.
4. Leaves are small, simple or compound. 4. Leaves are very large, simple; with sometimes
petiole as large as the depth of water.
5. The inflorescence may be cymose clusters, 5. The inflorescence is solitary terminal or solitary
racemose, or rarely solitary terminal. axillary.
6. Flowers usually small, actinomorphic or 6. Flowers usually large, long-peduncled, beautifully
zygomorphic. coloured, actinomorphic.
7. Sepals and petals vary in number from 3 to 7. Sepals usually 3 and petals usually many.
numerous; usually 5.
(Contd.)
Appendix 1 527
8. There is no transition of petals into stamens. 8. Petals (usually inner ones) frequently transform into
staminodes and stamens.
9. Several genera possess spurred sepals and 9. Sepals and petals are not spurred.
petals.
10. Gynoecium usually polycarpellary and 10. Carpels usually 3 and sometimes many, apocarpous
apocarpous. or syncarpous.
11. Placentation is basal or marginal. 11. Placentation is marginal or parietal.
12. Fruit an etaerio of achenes, or etaerio of 12. Fruit berry or follicle.
follicles, or berry, or capsule.
Table A.3 Ranunculaceae and Cruciferae (Brassicaceae)
Ranunculaceae Cruciferae (Brassicaceae)
1. Usually herbs, without a smelling watery juice. 1. Herbs with a smelling watery juice.
2. Leaves simple or compound. 2. Leaves simple and not compound.
3. Inflorescence may be cymose clusters, racemose, 3. Inflorescence usually corymbose raceme.
or flowers solitary terminal.
4. Sepals and petals may or may not be differentiated. 4. Sepals and petals always differentiated.
5. Sepals and petals vary in number from 3 to 5. Sepals and petals usually 4.
numerous, usually 5.
6. Sepals and petals are spurred in some genera 6. Sepals and petals never spurred.
(e.g. Delphinium).
7. Stamens 5 to many and never tetradynamous. 7. Stamens usually 6 and tetradynamous.
8. Gynoecium usually polycarpellary and apocarpous. 8. Gynoecium usually bicarpellary and syncarpous.
9. Ovary unilocular. 9. Ovary unilocular but becomes bilocular due to the
development of a false septum or replum.
10. Placentation is basal or marginal. 10. Placentation is parietal.
11. Stigma linear. 11. Stigma usually bifid.
12. Fruit an etaerio of achenes, or etaerio of follicles, 12. Fruit siliqua or silicula.
or berry, or capsule.
13. Seeds usually endospermic. 13. Seeds usually non-endospermic.
Table A.4 Ranunculaceae and Rosaceae
Ranunculaceae Rosaceae
1. Leaves compound, alternate but sometimes 1. Leaves compound, sometimes simple, alternate.
opposite.
2. Leaf bases mostly sheathing. 2. Stipulate.
3. Flowers usually regular, rarely irregular 3. Flowers regular, entomophilous but rarely
(Delphinium), entomophilous. anemophilous (Poterium).
4. Sepals variable in number and often petaloid. 4. Sepals usually 5.
5. Epicalyx absent. 5. Epicalyx often present.
6. Petals variable in number, often absent (Caltha). 6. Petals usually 5, sometimes absent (Alchemilla).
(Contd.)
528 Plant Taxonomy
7. Stamens numerous, spiral or cyclic. 7. Stamens numerous but only 4 in Alchemilla,

whorled.
8. Gynoecium of 3 to many carpels, separate. 8. Gynoecium of highly variable carpel number, ovary
free or adnate to hypanthium.
9. Fruit follicle, achene, and rarely berry (Actaea) 9. Fruit achene, aggregate, drupe (Prunus), pome
or capsule (Nigella). (Malus), follicle (Spiraea).
Table A.5 Papaveraceae and Fumariaceae
Papaveraceae Fumariaceae
1. Herbs usually with latex (Papaver, Argemone). 1. Herbs with watery juice but latex absent (Fumaria).
2. Usually the inflorescence is solitary terminal. 2. Usually the inflorescence is racemose.
3. Flowers actinomorphic. 3. Flowers zygomorphic.
4. Thalamus is always convex. 4. Thalamus is not always convex.
5. Petals rolled or crumpled in the bud. 5. Petals not always rolled or crumpled in the bud.
6. Usually the petals are unspurred. 6. Petals are usually spurred.
7. Stamens numerous. 7. Stamens are 6 and arranged in 2 groups.
8. Gynoecium bi- to polycarpellary. 8. Gynoecium is always bicarpellary.
Table A.6 Papaveraceae and Cruciferae (Brassicaceae)
Papaveraceae Cruciferae (Brassicaceae)
1. Plants mostly herbs with latex. 1. Plants mostly herbs which usually contain a
pungent watery sap.
2. Inflorescence solitary terminal. 2. Inflorescence typically a raceme or corymb.
3. Flowers usually bi- to trimerous. 3. Flowers usually tetramerous.
4. Corolla not cruciform. 4. Corolla cruciform (i.e. petals arranged in a cross).
5. Stamens numerous. 5. Stamens usually 6, arranged in two whorls.
6. Stamens do not show tetradynamous condition. 6. Stamens show tetradynamous condition.
7. Gynoecium bi-to polycarpellary, syncarpous. 7. Gynoecium always bicarpellary, syncarpous.
8. Locule does not bear a false septum or replum. 8. Usually the locule develops a false septum or
replum.
9. Fruit usually a capsule. 9. Fruit usually a siliqua or silicula.
Table A.7 Capparidaceae and Cruciferae (Brassicaceae)
Capparidaceae Cruciferae (Brassicaceae)
1. Includes herbs, shrubs and trees. 1. Plants are mostly herbs, and only rarely small
shrubs.
2. Plants usually devoid of smelling watery juice. 2. Plants with a smelling watery juice.
3. Leaves usually stipulate. 3. Leaves usually exstipulate.
4. Leaves simple or palmately compound. 4. Leaves simple or pinnately compound.
5. Flowers bracteate. 5. Bracts are absent.
(Contd.)
Appendix 1 529
6. Androgynophore or only gynophore present in 6. Androgynophore or gynophore absent.

some genera.
7. Corolla is never cruciform. 7. Corolla is usually cruciform.
8. Stamens 4 to numerous but never tetradynamous. 8. Stamens usually 6 and tetradynamous.
9. False septum or replum absent. 9. False septum or replum present.
10. Fruit berry, capsule, drupe or nut. 10. Fruit usually siliqua or silicula.
Table A.8 Capparidaceae and Caryophyllaceae
Capparidaceae Caryophyllaceae
1. Includes herbs, shrubs, small or tall trees. 1. Plants are mostly annual or perennial herbs.
2. Inflorescence usually raceme or simple umbel. 2. Inflorescence usually dichasial cyme.
3. Flowers generally tetramerous. 3. Flowers generally pentamerous.
4. Sepals usually 4, and rarely 5. 4. Sepals usually 5, and rarely 4.
5. Petals usually 4 and free. 5. Petals usually 5 and free.
6. Stamens are not obdiplostemonous. 6. Stamens are obdiplostemonous.
7. Androphore and gynophore present in 7. Androphore and gynophore absent.
several genera.
Table A.9 Malvaceae and Tiliaceae
Malvaceae Tiliaceae
1. Herbs or shrubs, rarely trees. 1. Usually trees or shrubs, rarely herbs.

2. Flowers always bisexual. 2. Flowers bisexual as well as unisexual.
3. Epicalyx usually present. 3. Epicalyx is not often present.
4. Sepals 5, free or basally connate, usually persistent. 4. Sepals 3–5, free or basally connate, caducous.
5. Stamens numerous, usually monadelphous. 5. Stamens numerous, usually polyadelphous.
6. Anthers monothecous, reniform, with spiny 6. Anthers dithecous, pollen grains smooth.
pollen grains.
7. Filaments of anthers fuse to form a staminal 7. Filaments do not form the staminal
column around the ovary. column around the ovary.
8. Seeds with curved embryo and scanty or 8. Seeds with straight embryo, and scanty
no endosperm. or copious endosperm.
9. Phloem fibres are not very strong. 9. Phloem fibres are very strong.
Table A.10 Oxalidaceae and Geraniaceae
Oxalidaceae Geraniaceae
1. Predominantly herbs, often shrubs or trees. 1. Predominantly herbs, rarely shrubs.

2. Leaves usually palmately compound, or 2. Leaves usually simple or sometimes compound,
pinnately compound, exstipulate. stipulate.
3. Leaves with a sour taste due to the presence 3. Oxalic acid absent.
of oxalic acid.
(Contd.)
530 Plant Taxonomy
4. Flowers actinomorphic. 4. Flowers actinomorphic but some are zygomorphic.

5. Sepals never spurred. 5. Sepals spurred in zygomorphic flowers.
6. Styles usually free. 6. Styles usually united.
7. Capsular fruit not dehiscing into 3–5 mericarps. 7. Capsular fruit dehiscing into 3–5 mericarps.
8. Seeds with straight embryo and fleshy 8. Seeds usually with curved embryo and scanty or no
endosperm. endosperm.
Table A.11 Rutaceae and Meliaceae
Rutaceae Meliaceae
1. Leaves are gland-dotted or contain translucent, 1. Leaves are not gland-dotted.

very clear dots.
2. Stamens obdiplostemonous. 2. Stamens not obdiplostemonous.
3. Filaments of anthers usually do not unite to form 3. Filaments unite to form a columnar tube.
a columnar tube.
4. Some genera (Citrus) show polyadelphous 4. Polyadelphous condition is very rare or absent;
condition. monadelphous condition is present.
5. A nectariferous disc present below the ovary. 5. Nectariferous disc usually present between
androecium and gynoecium.
6. Fruit usually a hesperidium. 6. Fruit is never hesperidium.
7. Seeds are not winged. 7. Seeds are often winged.
8. Polyembryony is very common. 8. Polyembryony is not of common occurrence.
Table A.12 Rutaceae and Anacardiaceae
Rutaceae Anacardiaceae
1. Leaves are gland-dotted or contain translucent, 1. Leaves are not gland-dotted and not even aromatic
very clear-dots; aromatic and strong-smelling. and strong-smelling.
2. Inflorescence cymose. 2. Inflorescence axillary or terminal panicles.
3. Flowers are hypogynous, mostly pentamerous. 3. Flowers hypogynous, rarely perigynous or
epigynous; pentamerous.
4. Cup-shaped nectariferous disc present between 4. Cup-shaped nectariferous disc is
ovary and stamens. extrastaminal or intrastaminal.
5. Sepals and petals mostly 4 or 5. 5. Sepals and petals mostly 5, rarely 3 to 7.
6. Stamens 8 to 10, obdiplostemonous; in some 6. Stamens usually 10, sometimes only 1 stamen and
members show polyadelphous condition 4 (Mangifera) or 6–9 staminodes (Anacardium).
(e.g. Citrus).
7. Gynoecium of 4 or 5 carpels, syncarpous; 7. Gynoecium of 1 to 5 fused carpels to form a
ovary tetra- to pentalocular, unilocular or multilocular ovary.
8. Fruit usually a hesperidium. 8. Fruit usually a drupe.
9. Polyembryony is very common. 9. Polyembryony is not very common.
Appendix 1 531
Table A.13 Sapindaceae and Anacardiaceae
Sapindaceae Anacardiaceae
1. Trees, shrubs or climbers, without any irritant or 1. Trees, shrubs or climbers having irritant or
acrid juice. acrid juice.
2. Resin ducts absent. 2. Resin ducts present.
3. Leaves usually pinnately compound. 3. Leaves simple or pinnately compound.
4. Flowers actinomorphic as well as zygomorphic. 4. Flowers mostly actinomorphic.
5. Sepals 4–5, poly- or gamosepalous. 5. Sepals 3–7 but usually 5, free or basally connate.
6. An annular disc is present, usually extrastaminal. 6. The cupular nectariferous disc is extrastaminal or
intrastaminal.
7. Gynoecium tricarpellary, syncarpous, trilocular. 7. Gynoecium of 1 to 5 fused carpels, unilocular or
multilocular.
8. Fruit drupe, berry, nut, capsule or samara. 8. Fruit usually a drupe.
Table A.14 Mimosoideae, Caesalpinioideae and Papilionoideae
Mimosoideae Caesalpinioideae Papilionoideae

1. Predominantly trees and 1. Mostly trees and shrubs, 1. Predominantly herbs, but
shrubs, rarely herbs. sometimes herbs. shrubs, trees and climbers
are also common.
2. Leaves mainly bipinnately 2. Leaves paripinnate, and 2. Leaves imparipinnate,
compound, and never never simple. sometimes simple.
simple.
3. Flowers actinomorphic. 3. Flowers zygomorphic. 3. Flowers zygomorphic.
4. Corolla aestivation valvate 4. Corolla aestivation asscending- 4. Corolla aestivation de-
or rarely imbricate. imbricate; posterior petal innermost. scending-imbricate; post-
erior petal outermost.
5. Petals 5, fused or free. 5. Petals 5, polypetalous. 5. Petals 5, unequal, 2
anterior petals basally
fused.
6. Stamens 10 or more. 6. Stamens usually 10, free or 6. Stamens usually 10 and
monadelphous, often a few stamens diadelphous i.e. (9) + 1,
reduce to staminodes. or monadelphous i.e. (10).
Table A.15 Rosaceae and Myrtaceae
Rosaceae Myrtaceae
1. Perennial herbs, shrubs or trees. 1. Usually large trees or shrubs.

2. Leaves stipulate, alternate or rarely opposite. 2. Leaves exstipulate, usually opposite.
3. Inflorescence highly variable from solitary 3. Inflorescence usually cymose.
flowers to racemose and cymose clusters.
4. Often epicalyx is present in several genera 4. Epicalyx is absent.
(e.g. Fragaria).
5. Sepals 5, free or basally connate. 5. Sepals 4–5, free or united.
(Contd.)
532 Plant Taxonomy
6. Petals 5, usually attached to the rim of 6. Petals 4–5, free or united.

hypanthium; sometimes numerous (e.g. Rosa).
7. Stamens numerous, usually in multiple of 5. 7. Stamens numerous with their anther connectives
gland-tipped.
8. Ovary superior to inferior with intermediate 8. Ovary inferior or semi-inferior.
stages.
9. Placentation is marginal or axile. 9. Placentation is axile or rarely parietal.
Table A.16 Rosaceae and Cucurbitaceae
Rosaceae Cucurbitaceae
1. Perennial herbs, shrubs or trees. 1. Mostly climbing or prostrate, tendril-bearing herbs;

shrubs and trees very rare.
2. Leaves stipulate. 2. Leaves exstipulate.
3. Flowers usually bisexual and only rarely 3. Flowers usually unisexual, and only rarely bisexual
unisexual (e.g. Pygeum). (e.g Schizopepon).
4. Epicalyx often present in several genera 4. Epicalyx absent.
(e.g. Fragaria, Geum).
5. Androecium usually made of numerous distinct 5. Androecium usually made of 5 stamens, usually
stamens in whorls of 5. more or less united into a column.
6. Usually a honey-secreting disc is present in 6. Disc is usually not present.
between stamens and carpels.
7. A transition between petals and stamens is 7. Such a transition is not seen.
usually seen as in Rosa.
8. Gynoecium usually varies from monocarpellary 8. Gynoecium usually tricarpellary.
to polycarpellary.
9. Ovary superior to inferior with intermediate 9. Usually the ovary is inferior.
stages.
10. Fruit of various types varying from drupe, 10. Fruit usually pepo or capsule.
pome, or follicle.
Table A.17 Rosaceae and Saxifragaceae
Rosaceae Saxifragaceae
1. Mostly trees or shrubs, and only rarely herbs. 1. Mostly herbs or shrubs, and only sometimes small
trees.
2. Leaves stipulate and not deciduous. 2. Leaves exstipulate and usually deciduous.
3. Inflorescence highly variable from solitary 3. Inflorescence usually cymose, and only sometimes
flowers to racemose and cymose. racemose to paniculate.
4. Epicalyx often present. 4. Epicalyx absent.
5. Sepals usually 5, free or united. 5. Sepals 4 or 5, often gamosepalous.
6. Petals usually 5 or even absent (e.g. Alchemilla). 6. Petals 4 or 5, sometimes connate or absent.
7. Stamens numerous, or 2-, 3-, or 4-times as many 7. Stamens usually twice as many as petals, in
as sepals, and whorled. 2 whorls, obdiplostemonous.
(Contd.)
Appendix 1 533
8. Ovary superior, sometimes inferior (Malus). 8. Ovary superior to inferior.

9. Carpels 1 to many, usually free. 9. Carpels usually 2, basally connate.
10. Fruit of various types from drupe (Prunus), 10. Fruit capsule or berry.
pome (Pyrus), follicle (Spiraea) to etaerio of
achenes (e.g. Rosa).
Table A.18 Umbelliferae and Rubiaceae
Umbelliferae (Apiaceae) Rubiaceae
1. Mostly herbs, and only a few are shrubs. 1. Mostly trees, or shrubs, and only some are herbs.
2. Mostly aromatic and contain essential oil or 2. Mostly do not contain essential oil or oleoresin.
oleoresin.
3. Leaves generally pinnately compound or 3. Leaves mostly simple.
decompound, and much divided; rarely simple.
4. Leaves exstipulate. 4. Leaves stipulate.
5. Inflorescence a simple or compound umbel. 5. Inflorescence basically a dichasial cyme.
6. Sepals 5, free or united, very small. 6. Sepals 4 or 5, free or basally united.
7. Petals 5, polypetalous. 7. Petals 4 or 5, gamopetalous.
8. Stamens 5, free and alternipetalous. 8. Stamens 4 or 5, epipetalous.
9. Styles often with a thickened or swollen base 9. Stylopodium absent; an epigynous disc often
called stylopodium. present.
10. Fruit a dry schizocarp called cremocarp, 10. Fruit a capsule (Cinchona), or berry (Mussaenda)
splitting into two mericarps. or schizocarp (Galium).
Table A.19 Umbelliferae (Apiaceae) and Compositae (Asteraceae)
Umbelliferae (Apiaceae) Compositae (Asteraceae)
1. Mostly aromatic herbs containing the essential 1. Herbs, shrubs, vines or rarely trees but not
oil or oleoresin in all organs. aromatic.
2. Leaves generally pinnately compound or 2. Leaves mostly simple, and only rarely truly
decompound, and much divided, rarely simple. compound.
3. Inflorescence simple or compound umble. 3. Inflorescence is a head or capitulum.
4. Disc florets and ray florets absent. 4. Disc florets and ray florets present.
5. Flowers are mostly actinomorphic. 5. Ray florets are usually zygomorphic.
6. Sepals are 5, free or fused, and never 6. Calyx represented by pappus.
represented by pappus.
7. Corolla of 5, free petals. 7. Corolla of 5, fused petals.
8. Stamens are 5, free, and never syngenesious. 8. Stamens 5, epipetalous, and syngenesious.
9. Ovary bilocular, and placentation axile. 9. Ovary unilocular, and placentation basal.
10. Stylopodium present. 10. Stylopodium absent.
11. Fruit is a cremocarp. 11. Fruit is a cypsela.
12. Seed with a minute embryo and copious 12. Seed with a large embryo and no endosperm.
endosperm.
534 Plant Taxonomy
Table A.20 Rubiaceae and Compositae (Asteraceae)
Rubiaceae Compositae (Asteraceae)
1. Mostly trees or shrubs, and only some are herbs. 1. Mostly herbs, only some are shrubs and only
rarely trees.
2. Leaves simple and stipulate. 2. Leaves simple and exstipulate.
3. Inflorescence basically a dichasial cyme. 3. Inflorescence basically a head or capitulum.
4. Disc florets and ray florets absent. 4. Disc florets and ray florets present.
5. Calyx made up of 4 or 5 sepals, polysepalous, 5. Calyx represented by pappus of bristles, awns or
well-developed. scales, or even absent.
6. Petals 4 or 5, gamopetalous. 6. Petals 5, gamopetalous; represented by 3 basic
types viz. (i) 5-lobed, tubular, (ii) ligulate,
and (iii) bilabiate.
7. Stamens 4 or 5, epipetalous and never 7. Stamens 5, epipetalous and syngenesious.
syngenesious.
8. Ovary inferior, bilocular. 8. Ovary inferior, unilocular.
9. Placentation axile. 9. Placentation basal.
10. Fruit capsule, berry or schizocarp. 10. Fruit a cypsela.
Table A.21 Asclepiadaceae and Apocynaceae
Asclepiadaceae Apocynaceae
1. Mostly perennial herbs, sometimes shrubs, 1. Mostly trees and shrubs, only some are herbs.
and rarely trees.
2. Stamens free or united to form a tube. 2. Stamens usually free.
3. Staminal corona present. 3. Corona absent.
4. Pollinia present. 4. Pollinia absent.
5. Translators and corpusculum present. 5. Translators and corpusculum absent.
6. Gynostegium or gynandrium present. 6. Gynostegium or gynandrium absent.
7. Nectariferous disc is absent. 7. Nectariferous disc is often present around or at the
base of the gynoecium.
8. Fruit a pair of follicles. 8. Fruit usually follicle, but sometimes a berry, drupe,
or capsule.
Table A.22 Convolvulaceae and Solanaceae
Convolvulaceae Solanaceae
1. Herbs, shrubs, or climbing or twining vines; with 1. Herbs or often shrubs but rarely vines; without
milky sap. milky sap.
2. Bicollateral bundles in stem absent. 2. Vascular bundles are often bicollateral in stem.
3. Calyx of 5, usually free, persistent sepals. 3. Calyx of 5, usually fused, persistent sepals.
4. Bracts often large and showy, sometimes 4. Bracts, if present, are not usually showy.
forming involucre.
(Contd.)
Appendix 1 535
5. Corolla often infundibuliform, tubular, 5. Corolla tubular to rotate, infundibuliform or bell-

salverform or campanulate; induplicately valvate shaped; rarely bilipped and zygomorphic
or twisted. (Schizanthus).
6. Anthers not connivent. 6. Anthers usually connivent.
7. Gynoecium bicarpellary syncarpous, but carpels 7. Gynoecium bicarpellary, syncarpous, but carpels
not obliquely placed. obliquely placed in the flower.
8. Placenta not swollen. 8. Placenta swollen.
Table A.23 Labiatae, Boraginaceae and Scrophulariaceae
Labiatae Boraginaceae Scrophulariaceae
1. Leaves opposite or whorled. 1. Leaves alternate. 1. Leaves alternate,

sometimes opposite.
2. Flowers zygomorphic, bilabiate. 2. Flowers actinomorphic. 2. Flowers zygomorphic.
3. Calyx of 5 united sepals, 3. Calyx of 5 united sepals, 3. Calyx of 5 united sepals.
sometimes 2-lipped. sometimes only basally fused
(Lithospermum).
4. Stamens usually 4, di- 4. Stamens 5, epipetalous. 4. Stamens usually 4,
dynamous, epipetalous, didynamous, sometimes
sometimes only 2 stamens. 2, rarely 5, epipetalous;
staminodes present.
5. Gynoecium with gynobasic 5. Gynoecium with 4-lobed ovary, 5. Gynoecium 2-carpellate.
style, 4-lobed ovary, bicarpellary.
bicarpellary.
6. Fruit 4 achenes (nutlets) 6. Fruit same as in Labiatae. 6. Fruit usually a capsule,
separating at maturity, sometimes a drupe or
sometimes a drupe. berry.
Table A.24 Acanthaceae, Labiatae and Verbenaceae
Acanthaceae Labiatae Verbenaceae
1. Mostly herbs or shrubs, rarely 1. Aromatic herbs or shrubs; with 1. Mostly shrubs or trees;
trees, not aromatic. square or quadrangular stem. stem often quadrangular;
plants not usually
aromatic.
2. Leaves simple, opposite 2. Leaves simple, opposite or 2. Leaves simple, opposite or
decussate, exstipulate. whorled, exstipulate; rarely whorled, exstipulate; some-
compound. times pinnately (Peronema)
or palmately (Vitex)
compound.
3. Inflorescence usually a 3. Inflorescence usually a 3. Inflorescence racemose
dichasial cyme. verticillaster. raceme, umbel, or spike.
4. Calyx of 5 united or free 4. Calyx of 5 united sepals, 4. Calyx of 5 united,
sepals. usually bilipped (1/4 in Ocimum persistent sepals.
and 3/2 in Salvia).
(Contd.)
536 Plant Taxonomy
5. Corolla of 5 united petals, 5. Corolla of 5 united petals, 5. Corolla of 5 united petals,

bilipped (2/3) or bilabiate bilipped (4/1 in Ocimum and 2/3 often bilipped.
personate. in Salvia, Leucas).
6. Stamens usually 4, didynamous, 6. Stamen usually 4, didynamous, 6. Stamens 4, didynamous,
epipetalous, rarely only 2 or 5 epipetalous, sometimes only epipetalous.
stamens. 2 stamens.
7. Style narrow and long, ovary 7. Style gynobasic, ovary 4-lobed. 7. Style single, terminal,
bilobed. ovary not lobed.
8. Fruit usually a capsule, rarely 8. Fruit schizocarp of 1–4 nutlets, 8. Fruit generally a drupe;
drupe. rarely drupe. rarely capsule, berry or
schizocarp.
9. Seeds with jaculators. 9. Seeds without any jaculator. 9. Seeds without any
jaculator.
Table A.25 Chenopodiaceae and Amaranthaceae
Chenopodiaceae Amaranthaceae
1. Predominantly halophytic or xerophytic annual or 1. Mostly annual or perennial, non-halophytic weedy

perennial herbs; only a few shrubs and rarely herbs; rarely shrubs.
small trees.
2. Leaves and stem mealy to touch. 2. Leaves and stem non-mealy to touch.
3. Inflorescence is not very dense or congested. 3. Inflorescence is usually very dense or congested.
4. Flowers monochlamydous, small and greenish. 4. Flowers monochlamydous, small with dry bracts.
5. Scarious bracts absent. 5. Usually scarious bracts present.
6. Tepals 5, usually persistent in fruit. 6. Tepals 5, dry or membranous, usually not persistent
in fruit.
7. Stamens 5, usually fee and not connate. 7. Stamens 5; usually connate filaments, at least at the
base.
8. Usually staminodes are not found. 8. Often with equal number of staminodes alternating
with the anthers (e.g. Celosia, Achyranthes).
Table A.26 Amaranthaceae and Polygonaceae
Amaranthaceae Polygonaceae
1. Mostly weedy herbs, rarely shrubs or climbers 1. Mostly weedy herbs with swollen node, a few
(Deeringia amaranthoides). climbers and only rarely trees (Triplaris).
2. Leaves exstipulate. 2. Leaves stipulate, and stipules are ochreate.
3. Inflorescence dense or congested, axillary or 3. Inflorescence primarily racemose but sometimes
terminal spikes, heads, or racemes, or flowers cymose, or clusters of panicled raceme (Rumex) or
solitary. cymose umbels (Eriogonum).
4. Perianth lobes usually 5, free or basally 4. Perianth lobes usually 3–6, free or basally connate.
connate.
5. Androecium usually of 5 stamens, opposite 5. Stamens usually 6–9 in two series.
tepals; often monadelphous (Achyranthes),
often with equal number of staminodes.
(Contd.)
Appendix 1 537
6. Gynoecium bi- to tricarpellary, syncarpous; 6. Gynoecium usually tricarpellary, syncarpous; ovary

ovary superior, unilocular. superior.
7. Ovule pendulous or campylotropous. 7. Ovule orthotropous.
8. Styles 1 to 3; stigma 1–3, capitate. 8. Style 1; stigma 2–4.
9. Fruit usually a utricle (Achyranthes) or 9. Fruit usually a triangular achene or nut
nutlet (Digera) or capsule (Celosia). (Polygonum).
Table A.27 Euphorbiaceae and Moraceae
Euphorbiaceae Moraceae
1. Herbs, shrubs or trees with milky latex; some 1. Mostly trees or shrubs, rarely herbs; contain
are xerophytes or cactus-like. milky juice.
2. Stem and leaves usually lacking stinging hairs. 2. Stinging hairs usually present.
3. Leaves stipulate, stipules sometimes in the form 3. Leaves often with 2 caducous stipules.
of hairs, glands, or thorns.
4. Several genera bear caducous or highly reduced 4. Leaves are well-developed in all members.
leaves.
5. Leaves of several genera contain extrafloral 5. Extrafloral nectaries absent.
nectaries.
6. Characteristic type of inflorescence is cyathium. 6. Syconium or hypanthodium is the characteristic
inflorescence of several genera.
7. Male flowers have one or more (sometimes up to 7. Usually the male flowers have only 4 stamens.
100, e.g. Croton) stamens.
8. Gynoecium is usually tricarpellary, syncarpous, 8. Gynoecium is usually bicarpellary, syncarpous,
superior, trilocular. superior to inferior and unilocular.
9. Placentation is usually axile. 9. Usual type of placentation is basal.
10. Fruit is usually a capsule or regma. 10. Usually a multiple fruit develops from the union
of fruits of several different flowers.
Table A.28 Musaceae and Zingiberaceae
Musaceae Zingiberaceae
1. Plants herbaceous but some appear like trees. 1. Plants herbaceous and none appears like tree.
2. Leaves large with long petiole and prominent 2. Leaves large or small; petiole and sheath not very
sheath. long; leaves may also be sessile.
3. Spathe usually conspicuous. 3. Bracteate.
4. Inflorescence usually a spadix or cincinnus. 4. Inflorescence usually a raceme or spike.
5. Perianth not distinguishable into calyx and 5. Calyx and corolla are distinct.
corolla.
6. Perianth lobes may be free or united. 6. Perianth lobes are usually fused.
7. Fertile stamens are 6 or 5. 7. Fertile stamen is only 1, and all others are modified
into staminodes.
538 Plant Taxonomy
Table A.29 Liliaceae and Iridaceae
Liliaceae Iridaceae
1. Leaves radical and not equitant. 1. Leaves equitant.

2. Inflorescence usually racemose, sometimes 2. Usually cymose inflorescence, sometimes flowers
umbellate (Allium) or solitary flowers (Tulipa). solitary (e.g. Sisyrinchium).
3. Flowers actinomorphic. 3. Flowers actinomorphic or zygomorphic (Gladiolus).
4. Perianth segments 6, often petaloid or sometimes 4. Perianth segments 6, often petaloid.
sepaloid.
5. Stamens 3 + 3, anthers usually introrse. 5. Stamens 3 + 3, anthers extrorse.
6. Ovary tricarpellary, syncarpous, superior. 6. Ovary tricarpellary, syncarpous, inferior.
7. Fruit loculicidal or septicidal capsule or berry. 7. Fruit loculicidal capsule.
Table A.30 Liliaceae and Amaryllidaceae
Liliaceae Amaryllidaceae
1. Storage organ usually rhizome, sometimes bulb, 1. Storage organ usually a bulb, and sometimes a
or roots are fleshy. rhizome.
2. Inflorescence usually racemose. 2. Inflorescence usually cymose.
3. Flowers usually actinomorphic. 3. Flowers actinomorphic or zygomorphic.
4. Tepals 3 + 3, free or united, petaloid or 4. Tepals 3 + 3, free or united, petaloid.
sometimes sepaloid.
5. Stamens usually 3 + 3. 5. Stamens usually 3 + 3 but sometimes staminodes
present.
6. Ovary superior. 6. Ovary inferior.
7. Fruit a loculicidal or septicidal capsule, or 7. Fruit usually a loculicidal capsule, rarely a berry.
a berry.
Table A.31 Juncaceae and Cyperaceae
Juncaceae Cyperaceae
1. Leaves usually in basal tufts, grass-like and 1. Leaves usually in basal tufts, grass like and
linear. 3-ranked.
2. Stem usually cylindrical, pith often present 2. Stem usually triangular and with solid pith.
but not solid.
3. Inflorescence cymose panicle or head. 3. Inflorescence is 1-many-flowered spikelets, arranged
variously in racemes, panicles, spikes, or umbels.
4. Perianth segments 6, in 2 whorls of 3 each; 4. Perianth segments often in the form of 6 bristles or
usually sepaloid. scales, or absent.
5. Stamens usually 6, in 2 whorls of 3 each, or 5. Stamens usually 1–3; only rarely 6 (Gahnia).
inner whorl absent.
6. Style simple, often with 3 brush-like stigmas. 6. Styles 2 or 3, or deeply divided into as many
branches as the number of carpels.
7. Fruit loculicidal capsule. 7. Fruit a triangular or biconvex achene or nutlet.
8. Seeds 3 to many in each fruit. 8. Seed 1 per fruit.
Appendix 1 539
Table A.32 Gramineae (Poaceae) and Juncaceae
Gramineae (Poaceae) Juncaceae
1. Stem jointed, usually cylindrical. 1. Stem usually without joints, cylindrical.

2. Pith is usually present only at the nodes. 2. Pith often present throughout.
3. Leaves basal, and, if cauline, 2-ranked, usually 3. Leaves mostly in basal tuft, usually linear, or terete.
linear to lanceolate.
4. Ligule usually present. 4. Ligule usually absent.
5. Basic unit of inflorescence is spikelet. 5. Inflorescence is usually cymose panicle or head.
6. Perianth is usually reduced into 2 or rarely 6. Perianth segments are generally 6, in 2 whorls of 3
3 lodicules. each, usually sepaloid.
7. Stamens usually 3 in each floret. 7. Stamens usually 6, in 2 whorls of 3 each, or inner
whorl is absent.
8. Styles usually 2, rarely 1 (Nardus) to 8. Style simple with 3 brush-like stigmas.
3 (Bambusa).
9. Fruit usually a caryopsis or rarely a nut, 9. Fruit a loculicidal capsule.
utricle (Sporobolus) or berry (some bamboos).
10. Seed usually 1 per fruit. 10. Seeds usually 3 to many in each fruit.
Table A.33 Cyperaceae and Gramineae (Poaceae)
Cyperaceae Gramineae (Poaceae)
1. Stem not jointed, usually triangular, with solid 1. Stem jointed, usually circular, with hollow
pith throughout. internodes; pith often present only at nodes.
2. Leaves 3-ranked or tristichous, leaf sheath 2. Leaves 2-ranked or distichous, leaf sheath
usually closed. usually open.
3. Leaves without ligule. 3. Ligule often present.
4. Bract, immediately subtending the flower, is 4. Bract, immediately subtending the flower, is usually
usually with odd number of nerves. 2-nerved.
5. Inflorescence various, from a simple spike to 5. Inflorescence spike, raceme or panicle.
a well-branched panicle.
6. Perianth represented often by 6 scales or bristles 6. Perianth usually in the form of 2 or 3 lodicules.
or absent, lodicules never present.
7. Stamens usually 1 to 3. 7. Stamens usually 3.
8. On reduction division, one microspore mother 8. On reduction division, one microspore mother
cell gives rise to 1 functional and 3 cell gives rise to a tetrad of 4 functional
non-functional microspores. microspores.
9. Style 1, often deeply divided into 2 or 9 Styles usually 2.
3 branches.
10. Fruit usually trigonous or biconvex achene 10. Fruit caryopsis.
or nut, and never a caryopsis.
11. Embryo usually embedded in the endosperm. 11. Embryo usually lateral to the endosperm.
EXAMINATION TOOLS MAJOR CHARACTERS OF DISCUSSED FAMILIES
540
Leaves Flowers Sepals Petals Stamens Ovary Carpels
Discussed
Major Distinguishing
Families of
Character/Characters
DICOTYLEDONS
Alternate
Opposite
Simple
Compound
Stipule
Actinomorphic
Zygomorphic
Bisexual
Unisexual
Free
United
Free
United
Many
Few
Superior
Inferior
Free
United
1. Magnoliaceae * * * * * + * * * * * + Deciduous stipules; perianth
lobes petaloid.
2. Annonaceae * * + * * * * * * * Leaves exstipulate, 2-ranked;
perianth 3 + 3 + 3, gynoecium
polycarpellary.
3. Ranunculaceae * + + * + * + * + * * * * * + Stamens numerous, spirally
arranged; carpels numerous.
4. Nymphaeaceae * * * * * * * * + + Aquatic; leaves with long
petiole.
5. Papaveraceae * * * * * + * * * + * Sepals deciduous; parietal
placentation.
6. Fumariaceae * * * * * * * * * Zygomorphic; spurred petals.
7. Capparidaceae * * + * * * * * * * * Androphore and gynophore
present.
8. Brassicaceae * * * + * * * * * * Corolla cruciform; stamens
(Cruciferae) tetradynamous; fruit siliqua or
silicula.
9. Violaceae * + * * + + * * * * * * Corolla spurred; one stamen
spurred.
10. Caryophyllaceae + * * + * * + + + * * * * Flowers obdiplostemonous;
placentation free-central.
11. Portulacaceae + + * * * * * * + * * + * Stipules scarious; sepals 2,
Plant Taxonomy
persistent.
(Contd.)
12. Malvaceae * * * * * + + * * + * * Epicalyx present; stamens mon-
adelphous; petals twisted.
13. Sterculiaceae * * * * * * + * * * * Stamens in 2 whorls; gynoe-
cium pentacarpellary.
Appendix 1
14. Tiliaceae * * * * * + + * * + * * Mostly trees or shrubs; stamens

many, polyadelphous.
15. Bombacaceae * * + * * * * * + * * * Large-sized trees; stipules cadu-
cous; flowers large-sized.
16. Geraniaceae + + + + * * + * * + * * * * Stamens with filaments united
at the base; fruit beaked or
lobed.
17. Oxalidaceae * + * * * * + * + * * * Leaves usually palmately com-
pound; sour due to presence
of oxalic acid.
18. Rutaceae + + + * * + * + + + * + + * * + * Fruit hesperidium; translucent
dots on the leaves; cymose
inflorescence.
19. Meliaceae * * * * + + + * + * * * Leaves exstipulate; monadel-
phous stamens; disc present
between androecium and
gynoecium.
20. Rhamnaceae * + * * * * + * + * * * + * Shrubs or trees, armed with
spines.
21. Vitaceae * + + + * * + + * * + * * * Climbing herbs; tendrils
present.
22. Sapindaceae * * * + * + + + * * * * Petals glandular; leaves
pinnately-compound.
23. Anacardiaceae * + + * * * + + * * * * * Intrastaminal disc present; fruit
drupe.
24. Leguminosae * + * * + * * + * + + + + * * Fruit a legume.
25. Subfamily * * * * * * + + + + * + Aestivation valvate or imbri-
Mimoseae cate; flowers actinomorphic.
541
(Contd.)
26. Subfamily * * * * * + + * + * * + Flowers zygomorphic;
Caesalpineae corolla aestivation ascending 542
imbricate.
27. Subfamily * * * * * * * * * + Aestivation descending
Papilionaceae imbricate.
28. Rosaceae * + + + * * + * + + + * * + * + * + Stamens numerous; fruits of
various types.
29. Saxifragaceae * * * * + + * + * + + * Leaves deciduous, exstipulate;
ovary superior to inferior.
30. Combretaceae * + * * + * + * * * * * Often lianous; hypanthium
present.
31. Myrtaceae + * * * * + + + + * + * * Trees or shrubs; anther con-
nectives gland-tipped; ovary
inferior.
32. Lythraceae * * + * + * * * + * * * Corolla crumpled; ovary supe-
rior; epicalyx present.
33. Cucurbitaceae * * * + * + * + * * * * Fruit pepo; tendril-bearing
herbs.
34. Begoniaceae * * * + * * * + * + * * * Unisexual flowers; inferior
ovary.
35. Passifloraceae * * + * * * + + + + + * * * Climbing habit; uniflowered
peduncles; androgynophores
present.
36. Cactaceae * * + + * * * * * * Mostly prickly, spiny, fleshy,
succulents.
37. Umbelliferae * + * * * + + + * * * * Inflorescence simple or com-
(Apiaceae) pound umbel; inferior ovary.
38. Araliaceae * + * * * + + + + * * + * * Leaves alternate; flowers
actinomorphic.
39. Rubiaceae * * * * + * * * * * * Stamens alternipetalous; inferior
Plant Taxonomy
ovary.
(Contd.)
40. Caprifoliaceae * * + + + + * + * * * * * Calyx fused to the ovary;
multicarpellary.
41. Compositae * + * + + + + + * * * * Calyx usually a pappus; head
(Asteraceae) or capitulum.
Appendix 1
42. Campanulaceae * + * + + * * * * + * * Stamens epipetalous and

show connation of anthers or
filaments.
43. Ericaceae * + * * + * + + * + * * * + * Flowers urceolate; ovary tetra-
to multilocular.
44. Primulaceae + * * * + * * * * * * Stamens opposite the petals;
free-central placentation.
45. Plumbaginaceae * * * * * + * * * * 5-styled pistil; fruit dry,
1-seeded.
46. Sapotaceae * * * * + * * * * * Plants with milky latex; sta-
mens in 2-3 whorls.
47. Oleaceae + * + + * * + * * * * * Trees or shrubs with opposite
leaves; stamens 2.
48. Asclepiadaceae * * + * + * + + * * * * + Herbs with milky sap; corona,
pollinia, translators and
corpusculum present.
49. Apocynaceae + * * + * * * * * * * Stamens epipetalous; corona,
pollinia and corpusculum
absent.
50. Loganiaceae * * * * * * * * * * Simple, opposite, stipulate
leaves; internal phloem
present.
51. Boraginaceae * + * * + * + + * * * * Hispid herbs; inflorescence
helicoid cyme; style
gynobasic.
52. Convolvulaceae * * * * * + * * * * Corolla tubular,
infundibuliform of salverform.
53. Solanaceae * + * * + * * * * * * Carpels obliquely placed.
543
(Contd.)
54. Polemoniaceae + + + + * + * * * * * * Stamens attached at different
lengths on corolla tube; ovary 544
tricarpellary.
55. Scrophulariaceae + + * + + * * * * * * * Flowers zygomorphic; corolla
bilipped.
56. Bignoniaceae * * * * * * * * * Inflorescence cymose, flowers
zygomorphic.
57. Pedaliaceae + * * * * * * * * * Mostly herbs; stamens 4,
didynamous.
58. Acanthaceae * * * * + * * * * * Flowers zygomorphic; corolla
bilipped (2/3); seeds with
jaculators.
59. Lamiaceae * * * * * * * * * Inflorescence verticillaster;
(Labiatae) coralla bilipped (4/1 or 2/3);
style gynobasic.
60. Verbenaceae + * * + * * * * * * * Stem quadrangular; ovary not
lobed; style terminal.
61. Plantaginaceae * + * * * * * * * * Leaves with somewhat parallel
venation; petals membranous.
62. Chenopodiaceae * + * * * + + * ---------- * * * Perianth sepaloid; halophytic
herbs; fruit utricle.
63. Amaranthaceae + + * * * + + + ---------- * * * Mostly herbs; bracts dry, scari-
ous; stamens connate; perianth
sepaloid.
64. Polygonaceae * * * * * + ---------- + + * * * Tepals petaloid; ochreate
stipules.
65. Aristolochiaceae * * + + * * ---------- + + * * Sepals petaloid; petals absent.
66. Piperaceae * + * + * * + ---------- ---------- * * * Perianth absent; stamens 1-10;

embryo very minute.
67. Loranthaceae + * * * + + + + + + + * * * Aerial, parasitic habit; cup-
Plant Taxonomy
shaped receptacle.
(Contd.)
68. Euphorbiaceae * + * + * * * * + * + + * * * Milky latex present; flow-
ers unisexual; fruit capsule or
regma; inflorescence cyathium.
69. Urticaceae + + * + * * + + ---------- * * * Herbs with stinging hairs and
Appendix 1
watery sap; mostly sepaloid.

70. Moraceae * * * * * + + + + * + + * Leaves stipulate; flowers
uni-sexual; tepals usually 4.
71. Cannabinaceae + + + + * * * + + + + * * * Monocarpellary, calyx persis-
tent; unisexual.
72. Casuarinaceae + + + + * ---------- ---------- * * * Equisetum-like jointed stem;
minute scaly leaves; unisexual
flowers.
73. Salicaceae * + * * * * ---------- ---------- + * * * Deciduous dioecious trees or
shrubs; flowers in catkins; a
cup-like disc or gland present
in each flower.
MONOCOTYLEDONS
1. Orchidaceae * * * * + * + * + * * * Leaves 2-ranked; labellum,
pollinia and gynostegium
present.
2. Iridaceae ---------- * + + * + + + + * * * Leaves mostly basal and
equitant; herbs with bulbs,
corms or rhizome; trimerous.
3. Amaryllidaceae * * * + * + + + + * * * Inflorescence a long leafless
scape; bulbous or rhizomatous
herbs; stamens antiphyllous.
4. Bromeliaceae + * * + * + * + + * + * * Short-stemmed herbs; leaves
stiff; anthers versatile.
5. Cannaceae * * * * * * * * * Leaves very large with petiole
sheathing their base; inflores-
cence a 2-flowered cincinii.
545
(Contd.)
6. Musaceae * * * + * + + + + * * * Perennial giant herbs; leaves
546
very large; inflorescence spa-
dix covered by a spathe.
7. Zingiberaceae * * * * * + * * * * Aromatic herbs; only one
fertile stamen; ligule present at
the top of leaf sheath.
8. Liliaceae * + * + * * + + + + + * * * Herbs with bulbs, corms or
rhizome; perianth showy.
9. Commelinaceae * * * + * * * + * * * Herbs with succulent stem;
leaves with tubular sheath;
ovary superior.
10. Juncaceae * * * + * * * * * Leaves mostly basal; perianth
sepaloid.
11. Palmae + + * * + * + + + + * * + * Leaves usually in terminal clus-
(Arecaceae) ters and leave prominent scars
of leaf bases.
12. Typhaceae * * * * ---------- ---------- * * * Perianth often of hairs or scales;
fruit achene; monocarpellary.
13. Araceae * + + * + * + + + + * * + * Inflorescence spadix enveloped
by a spathe; aquatic or
terrestrial habit.
14. Alismataceae ---------- * * + + * * + * * * Marshy herbs with perennating
rhizomes and basal leaves.
15. Cyperaceae * * * + * + ---------- ---------- * * * Herbs with triangular stem;
leaves 3-ranked, bract gluma-
ceous; fruit achene or nut.
16. Gramineae * * * * + ---------- ---------- * * * Perianth represented by
(Poaceae) lodicules; ligule usually pres-
ent; leaves 2-ranked; fruit
caryopsis.
*, present; +, present only sometimes; -------, absent or represented by bristles or minute scales. In the families where symbols appear in both columns for contrasting
Plant Taxonomy
characters (+ * and * +); * is the typical condition but sometimes + is found; + +, both characters equally common. Columns left blank or without any symbol
means that this character is not found in the particular family.
APPENDIX 2
EXAMINATION TOOLS SELECTED MEDICINAL PLANTS AND
THEIR UTILITY: AT A GLANCE
Name of the Plant * Family Parts Used in Treatment of
1. Michelia champaca Magnoliaceae Flowers and fruits (kidney trouble, gonorrhoea).

2. Illicium verum Magnoliaceae Seeds (colic disorders).
3. Uraria narum Annonaceae Roots (urinary problems).
4. Aconitum napellus Ranunculaceae Roots (nerve sedative, rheumatism, heart palpitation,
gastritis).
5. Actaea spicata Ranunculaceae Roots (nerve sadative).
6. Anemone pulsatilla Ranunculaceae Plant extract (nervous disorders in women during
menstrual periods).
7. Coptis teeta Ranunculaceae Roots (brain and liver disorders).
8. Delphinium zatil Ranunculaceae Seeds, leaves and roots (jaundice, enlarged spleen).
9. Delphinium vestitum Ranunculaceae Seeds (cardiac disorders).
10. Helleborus niger Ranunculaceae Flowers (madness).
11. Nigella sativa Ranunculaceae Seeds (cough, asthma, fever).
12. Paeonia officinalis Ranunculaceae Tubers (colic and uterus disorders).
13. Ranunculus aquatilis Ranunculaceae Flowers (asthama and rheumatism).
14. Ranunculus ficaria Ranunculaceae Flowers and seeds, (piles).
15. Nelumbo nucifera Nymphaeaceae Flowers (cardiac tonic and liver disorders),
rhizome (piles).
16. Nymphaea nouchali Nymphaeaceae Rhizome (dysentery and diarrhoea).
17. Nymphaea stellata Nymphaeaceae Flowers juice (cardiac pain).
(Contd.)
* Plants arranged according to families discussed in the book.

548 Plant Taxonomy
18. Papaver somniferum Papaveraceae Milky latex of unripe fruits produce “opium” (known
for its sedative properties).
19. Argemone mexicana Papaveraceae Seeds (skin infections).
20. Sanguinaria canadensis Papaveraceae Rhizome and roots (emetic and dyspepsia i.e.
indigestion).
21. Hypecoum procumbens Fumariaceae Alkaloid protopine (high blood pressure).
22. Corydalis govaniana Fumariaceae Roots (syphilis).
23. Fumaria indica Fumariaceae Dried plants and seeds (fever and also as blood
purifier).
24. Cleome gynandra Capparidaceae Leaves and seeds (expelling roundworms).
25. Cleome religiosa Capparidaceae Bark (increase appetite; reduce bile secretion).
26. Brassica oleracea var. Brassicaceae Inflorescence (possess anticancer properties).
botrytis (Cauliflower)
27. Viola odorata (Banafsha) Violaceae Dried flowers (cough, influenza).
28. Hybanthus enneaspermus Violaceae Roots (urinary tract infections, gonorrhoea).
29. Dianthus sinensis Caryophyllaceae Flowers and seeds (gonorrhoea).
30. Spergula arvensis Caryophyllaceae Seeds (lungs tuberculosis).
31. Saponaria vaccaria Caryophyllaceae Plant extract (urinary bladder diseases).
32. Stellaria media Caryophyllaceae Leaves (inflammations of digestive and respiratory
tracts).
33. Stellaria semivestita Caryophyllaceae Entire plant (anticancerous).
34. Portulaca oleracea Portulacaceae Entire plant extract (ailments of kidney and urinary
bladder).
35. Hibiscus rosa-sinensis Malvaceae Roots (cough and cold).
36. Althaea rosea Malvaceae Roots (dysentry).
37. Gossypium sp. Malvaceae Bark (stops haemorrage after child birth in ladies).
38. Malva verticillata Malvaceae Roots (whooping cough).
39. Urena repanda Malvaceae Roots and bark (hydrophobia).
40. Helicteres isora Sterculiaceae Juice of roots (diabetes).
41. Corchorus capsularis Tiliaceae Dried leaves and roots decoction (dysentry and
and C. olitorius diarrhoea).
42. Grewia asiatica Tiliaceae Root bark (rheumatism).
43. Grewia tiliaefolia Tiliaceae Wood (emetic and as antidote of opium poisoning).
44. Triumfetta bartramia Tiliaceae Leaves, fruits and flowers (gonorrhoea).
45. Durio zibethinus Bombacaceae Root decoction (fever).
46. Geranium nepalense Geraniaceae Root and leaf decoction (kidney troubles).
47. Acranychia laurifolia Rutaceae Bark poultice (ulcers and sores).
48. Aegle marmelos Rutaceae Fruit pulp (mild laxative), roasted fruits (diarrhoea),
bark (intermittent fever).
49. Atalantia monophylla Rutaceae Oil from fruits (paralysis and rheumatism).
50. Barosma betulina Rutaceae Leaves (kidney diseases).
(Contd.)
Appendix 2 549
51. Cusparia febrifuga Rutaceae Bark (malaria).

52. Dictamnus albus Rutaceae Bark (nervous disorders and intermittent fever).
53. Feronia limonia Rutaceae Ripe fruits (cardiac tonic).
54. Peganum harmata Rutaceae Seeds (asthma, neuralgia and rheumatism).
55. Pilocarpus pinnatifolius Rutaceae Leaves (kidney troubles).
56. Toddalia asiatica Rutaceae Root bark (antimalarial).
57. Zanthoxylum alatum Rutaceae Seeds (cholera).
58. Azadirachta indica Meliaceae Seeds (skin diseases), bark (malaria), twigs (pyorrhea).
59. Melia azedarach Meliaceae Leaf juice (anthelmintic), seeds (rheumatism).
60. Aglaia odorata Meliaceae Fruits (leprosy).
61. Dysoxylum malabaricum Meliaceae Wood oil (eye and ear diseases).
62. Walsura piscidia Meliaceae Bark (stimulant, expectorant and skin diseases).
63. Rhamnus purshiana Rhamnaceae Bark (purgative).
64. Cayratia carnosa Vitaceae Leaves and roots (high fever).
65. Leea aequata Vitaceae Oil from the plant (tuberculosis).
66. Leea macrophylla Vitaceae Tubers (ringworms).
67. Acer indica Sapindaceae Fruits and seeds (rheumatism).
68. Caesalpinia bunduc Caesalpinioideae Seeds (diarrhoea, rheumatism).
69. Cassia fistula Caesalpinioideae Fruit pulp (purgative).
70. Cassia occidentalis Caesalpinioideae Leaves and seeds (skin infections).
71. Cassia sophera Caesalpinioideae Leaf decoction (acute bronchitis).
72. Crotalaria albida Papilionoideae Roots (purgative).
73. Glycyrrhiza glabra Papilionoideae Roots (cough and sore throat).
74. Krameria triandra Papilionoideae Roots (chronic diarrhoea).
75. Moghania strobilifera Papilionoideae Roots (induce sleep).
76. Psoralea corylifolia Papilionoideae Seeds (leucoderma and leprosy).
77. Teramnus labialis Papilionoideae Entire plant (tuberculosis).
78. Rosa sp. Rosaceae Petals yield “Gulkand” (tonsilitis, cough, cold; tonic).
79. Hagenia abyssinica Rosaceae Female flowers (source of “cusso”, an antihelmintic
drug).
80. Potentilla anserina Rosaceae Leaf decoction (arthritis and kidney stones).
81. Terminalia arjuna Combretaceae Bark (cardiac tonic).
82. Terminalia bellirica Combretaceae Fruits (“Bahera”, one of the three constituents of
Triphala, along with Hararh and “Amla”).
83. Terminalia chebula Combretaceae Fruits (“Hararh”, one of the three constituents of
Triphala).
84. Combretum roxburghii Combretaceae Leaves (malaria fever).
85. Syzygium aromaticum Myrtaceae Clove oil (indigestion, antiseptic, antispasmodic).
86. Eugenia jambolana Myrtaceae “Jamun” fruits (diabetes).
87. Ferula assafoetida and Umbelliferae Dried latex from roots (“Heeng”, stimulates respiratory,
F. narthax nervous and intestinal systems).
(Contd.)
550 Plant Taxonomy
88. Centella asiatica Umbelliferae “Brahmi Booti” (brain tonic, used in madness and
leprosy).
89. Ferula sumbule Umbelliferae Plant decoction (hysteria).
90. Panax ginseng Araliaceae Ginseng roots (stimulant and aphrodisiac).
91. Cinchona sp. (C. calisaya, Rubiaceae Bark (yield quinine, used in malaria).
C. officinalis)
92. Cephaelis ipecacuanha Rubiaceae Roots (amoebic dysentery).
93. Anthemis mobilis Compositae Dried capitula (dyspesia).
94. Artemisia cina Compositae Flower heads (drug “santonin” used in intestinal
worms).
95. Blumea balsamifera Compositae Leaves (curie excitement and insomnia).
96. Grindelia camporum Compositae Floral heads (bronchitis, whooping cough).
97. Inula helium Compositae Leaves (tuberculosis).
98. Sphaeranthus indicus Compositae Floral heads (stomach ache, piles).
99. Tanacetum vulgare Compositae Leaves (chronic ulcers, rheumatism).
100. Lobelia inflata Campanulaceae Dried leaves and tops (drug “lobelia” is used as
expectorant and emetic, and in bronchitis and asthma).
101. Anagallis arvensis Primulaceae Plant decoction (leprosy, gout, hydrophobia).
102. Achras sapota Sapotaceae Chicle gum (dental surgery).
103. Mimusops elengi Sapotaceae Seeds and dried fruits (piles).
104. Madhuca indica Sapotaceae Oil from seeds (skin diseases, rheumatism).
105. Nyctanthes arbortristis Oleaceae Leaves (fever, rheumatism).
106. Calotropis procera Asclepiadaceae Roots (cough).
107. Tylophora indica Asclepiadaceae Roots (bronchitis, asthma, whooping cough).
108. Rauvolfia serpentina Apocynaceae Dried roots (“Sarpagandha” is used in hypertension,
mental disorders and schizophrenia).
109. Thevetia peruviana Apocynaceae Seeds (rheumatism).
110. Alstonia scholaris Apocynaceae Bark (malaria and dysentery).
111. Catharanthus roseus Apocynaceae Plant extract (leukaemia)
112. Strychnos nux-vomica Loganiaceae Seeds (nervous disorders, paralysis).
113. Strychnos toxifera Loganiaceae Seeds (shock therapy, muscle relaxant, tetanus).
114. Lithospermum officinale Boraginaceae Seeds (urinary bladder diseases).
115. Exogonium purga Convolvulaceae Tuberous roots (purgative).
116. Nicotiana tabacum Solanaceae Dried leaves (sedative. antispasmodic).
117. Atropa belladona Solanaceae Roots (drug at tropine relieves spain; dilates pupil of
eyes).
118. Datura stramonium Solanaceae Leaves and flowers (drug stramonium used in asthma).
119. Mandragora autumnalis Solanaceae Roots (sedative and hypnotic).
120. Withania coagulans Solanaceae Fruits (asthma and liver troubles).
121. Withania somnifera Solanaceae Roots (the drug “asgandha” cures cough, rheumatism,
sexual weakness; promotes urination).
(Contd.)
Appendix 2 551
122. Bacopa monniera Scrophulariaceae Decoction of mature plants (epilepsy, insanity).

123. Digitalis purpurea Scrophulariaceae Dried leaves (congested heart failure).
124. Adhatoda vasica Acanthaceae Leaves (cough, asthma, bronchitis).
125. Hygrophila spinosa Acanthaceae Leaves and roots (jaundice, rheumatism).
126. Ocimum sanctum Labiatae Leaves (cough, cold, fever).
127. Mentha spicata Labiatae Leaves (indigestion, rheumatism).
128. Thymus vulgaris Labiatae Leaves (hookworm infection).
129. Chenopodium ambrosioides Chenopodiaceae Decoction of plant (hookworm infection).
130. Achyranthes aspera Amaranthaceae Inflorescence (antidote against snake and scorpion
bites).
131. Polygonum aviculare Polygonaceae Leaves and seeds (diabetes, rheumatism).
132. Rheum emodi and Polygonaceae Roots and rhizomes (drug “rhubarb” is used as laxative,
R. officinale purgative and stomach tonic).
133. Rumex acetosella Polygonaceae Leaves (tumours and cancer).
134. Polygonum glabrum Polygonaceae Seeds (leucorrhoea).
135. Emblica officinalis Euphorbiaceae Fruits (“Aamla”, rich in vitamin C; used in scurvy;
one of the three constituents of “Triphala”).
136. Ricinus communis Euphorbiaceae Seeds (purgative).
137. Jatropha gossypifolia Euphorbiaceae Leaves (eczema), root (leprosy).
138. Synadenium grantin Euphorbiaceae Entire plant (stimulates central nervous system).
139. Morus alba and M. nigra Moraceae Bark (purgative and vermifuge).
140. Antiaris toxicaria Moraceae Latex (cardiac stimulant).
141. Salix sp. Salicaceae Soft inner bark and root (“aspirin” is used for
headache).
142. Belamcanda chinensis Iridaceae Rhizome (tonsilitis).
143. Iris germanica Iridaceae Roots (gall bladder diseases).
144. Curculigo orchioides Amaryllidaceae Tubers (asthma, jaundice).
145. Curcuma longa Zingiberaceae Rhizome (“turmeric” is used as stomachic,
antiseptic, and blood purifier).
146. Zingiber officinale Zingiberaceae Rhizome (carminative, digestive stimulant).
147. Aloe barbadense Liliaceae Leaves (purgative, piles, inflammations).
and A. vera
148. Asparagus racemosus Liliaceae Roots (nervous and rheumatic complaints; skin
diseases).
149. Colchicum luteum Liliaceae Roots (gout, rheumatism).
150. Gloriosa superba Liliaceae Tubers (promoting labour pains in women).
151. Smilax glabra and Liliaceae Roots (veneral diseases).
S. ovatifolia
152. Veratrum viridae Liliaceae Roots (hypertension).
153. Cyperus scariosus Cyperaceae Tubers (stone trouble of kidney and urinary bladder).
154. Cyperus stoloniferous Cyperaceae Tubers (heart stimulant).
INDEX
Aadu 317 Adina 313, 350 Alligator weed 432 Anacardium 293
Aak 384 Adnate 199 Allium 101, 490, 493 Anagallis 369, 550
Aalu 400 Adonidia 502 Allolepis 524 Ananas 479, 480, 482
Aamla 323 Adonis 223 Almond 312 Anatomy In Relation To Taxonomy 98
Abelia 352 Adoxa 355 Alocasia 507, 509 Anchusa 393
Abelmoschus 260 Adoxaceae 121, 349 Aloe 9, 162, 490, 491, 492, 493, 494, Andaman redwood 309
Abir 490 Adrak 489 551 Andesia 497
Abortion hypothesis 145 Adventitious Root 177 Aloeaceae 494 Androecium 65
Abroma 262, 289 Aechmea 480 Aloocha 317 Androgynophore 194, 240, 276
Abrus 309 Aegilops 105 Aloysia 425 Androphore 194, 240
Abutilon 257, 105, 222, 223 Aegle 276, 548 Alpinia 488, 489, 490 Andropogon 523
Acacia 299, 75 Aegopodium 345 Alsodeia 247 Androsace 369
Acaena 312 Aerial 177 Alstonia 385, 220, 550 Aneilema 494, 495
Acalypha 446, 221, 222 Aeschynomene 310 Alstroemeria 478, 479 Anemone 70, 384, 406, 407, 348, 547
Acanthaceae 25, 38, 535, 544, 414 Aesculus 21 Alternanthera 432 Anemone, 9
Acanthaceae, Scrophulariaceae 34 Aestivation 196 Alternate 186 Anemonella 105
Acantholimon 374 Agapanthus 491 Alternipetalous 199 Anemopaegma 409
Acanthopanax 348 Agarbattis 517 Althaea 257, 323, 548 Anethum 343
Acanthosicyos 330 Agatea 247 Althaea, Malva 258 Aneuploidy 104
Acanthus 414, 28 Agavaceae 100, 479, 504 Alu-balu 317 Angelica 345
Acaulescent 179 Agave 105, 477, 478, 479 Alubukhara 317 Angelonia 406
Acer 22, 179, 377, 549 Ageratum 355 Alyssum 97 Angiospermae 13, 27, 37
Aceraceae 297 Agglutination 134 Amaranthaceae 25, 536, 544 Angiosperm phylogeny 139, 140
Achene 206, 376 Agglutinins 134 Amaranthaceae1 428 Angiosperms 12, 72
Achillea 356 Agglutinogens 134 Amaranthus 432 Angular 179
Achlamydosporeae 26, 101, 34 Aglaia 282, 549 Amar Bel 397 Animalia 47
Achras 375, 433, 434, 550 Agrimonia 312 Amaryllidaceae 26, 477, 479, 482, 493, Anjeer 455
Achyranthes 29, 279, 432, 435, 536, Agropyron 518 538, 545 Annona 218
537, 551 Agrostis 518, 523 Amaryllidaceae1 469 Annonaceae 24, 217, 526, 540
Acioa 314, 42 Aizoaceae 123, 430 Amaryllis 477, 478 Annonidae 13, 320
Aclimandra 214, 42 Ajwain 343 Amborella 101 Annonopsida 13, 310, 469
Acnidia 435 Akash-Bel 397 Amborellaceae 101 Annual 176
Acokanthera 387 Akash Neem 411 Ambrosia 359 Annulated 177
Aconite 223 Akee 292 Ambulia 406 Anogeissus 321
Aconitum 136, 125, 547 Albizzia 299 Amentiferae 43, 352 Anshphal 292
Acorus 507, 508, 509 Alcea 257 American Code 81 Anthemis 356, 550
Acotyledones 22 Alchemilla 312, 527 Amherstia 304 Anthericum 97
Acranychia 280, 548 Aleurites 450 Amino acids 125 Anthocephalus 350
Actaea 136, 43, 547 Alexandra 430 Amla 449 Anthocyanins 121
Actinomorphic 166, 194 Alfalfa 306, 308 Ammodendron 125 Anthophore 194
Actinostemma 330 Algae 22, 27 Amomum 488, 489 Anthurium 507, 509
Acuminate 182 Alisma 510, 511, 512, 513 Amoora 282 Anthyllis 97
Acute 182 Alismaceae 26, 294 Amorphophallus 507, 509 Antiaris 456, 551
Adansonia 269 Alismaceae1 470 Amorphosporophyta 108 Antibody 134
Adansonian classifications 115 Alismalidae 38 Ampelocissus 286, 359 Antigen 134
Adenanthera 302 Alismataceae 28, 509, 513, 546 Ampelopsis 288 Antigonon 250, 359, 348, 355
Adenia 336 Alismatales 31 Amrood 325 Antirrhinum 405
Adenium 385 Alismatidae 39 Amsinckia 392 Apetalous 197
Adenocalymna 411 Alismatiflorae 41 Anacampseros 254 Aphania 290
Adenocarpus 125 Alkaloids 121, 124 Anacardiaceae 25, 293, 530, 531, 541 Aphanic species 50
Adhatoda 52, 105, 414, 551 Allamanda 385 Aphelandra 414
554 Index
Apiaceae 52, 342, 533, 542 Ashok tree 304 Basak 415 Boehmeria 451
Apiculate 182 Asimina 219 Basal 205 Boenninghausenia 277
Apium 343 Asiphonogama 28 Basella 429 Bombacaceae 29, 268, 541
Apluda 523 Asparagus 8, 490, 491, 492, 493 Basellaceae 123 Bombaceae 109
Apocarpae 26 Asperula 137 Basifixed 200 Bombax 269, 342
Apocarpeae 470 Asphodelus 490, 491, 494 Basionym 87 Bonplandia 404
Apocarpous 202 Aspidosperma 385 Bathua 430 Borage 393
Apocynaceae 25, 534, 543 Aspirin 464 Batidaceae 29 Boraginaceae 30, 426, 535, 543
Apocynaceae 384 Assimilatory 177 Batidales 29 Borago 393
Apocynaceae, Asclepiadaceae 34 Aster 104 Bauhinia 108 Borassus 500, 501, 502, 503
Apocynum 119 Asteracantha 415 Bean 309 Botanical capital of the world 163
Apopetalous 197 Asteraceae 12, 355, 533, 534 Beaumontia 386 Botanical gardens 158
Aporosa 450 Asterales 25 Bedina 352 Botanical library 148
Apostasia 472 Asteridae 38 Begonia 333 Botanical names 80, 89
Appanomixis 281 Astilbe 318 Begoniaceae 25, 257, 440, 542 Botanical Survey of India, 16, 17
Apple 316 Astragalus 306 Begoniella 334 Bottle brush 326
Apricot 312 Astrantia 342 Belamcanda 474, 475, 476, 551 Bottle gourd 330
Aquatic 176 Atalantia 277, 548 Belladona 400 Bottle palm 502
Aquatic geophytes 145 Atom 128 Beloperone 415 Bouea 293
Aquifoliaceae 85 Atriplex 126 Benincasa 331 Bougueria 426
Aquilegia 70 Atropa 399, 550 Bennettitalean Theory 142 Bourdon wine 503
Arabis 243 Aurone 122 Ber 286 Brachychilum 490
Araceae 26, 504, 507, 509, 546 Australian Acacia 302 Berberidaceae 109, 276 Brachychiton 265
Araceae1 470 Authority 80 Berberidales 233 Brachyelytrum 524
Arachis 306 Autonym 87 Berberis 21 Bracket or parallel key 71
Arales 31 Avena 518, 520, 522, 524 Berchemia 285, 430 Bract 169, 184
Aralia 346 Averrhoa 274 Bergamot oil 280 Bracteate 195
Araliaceae 98, 542 Averrhoaceae 276 Bergenia 320 Bracteolate 195
Arandi 450 Avicennia 423 Berry 206 Bracteole 169, 184
Arborae 20 Axile 205 Berrya 267 Brahmi Booti 343
Arborescent 179 Axis 169 Besseyan Principles 30, 139 Brasenia 228
Arbutus 366, 440 Axonopus 523 Bessey’s cactus 31 Brassica 104, 548
Archegoniatae 28 Azadirachta 281, 549 Beta 429, 243, 244 Brassicaceae 34, 243, 527, 528, 540
Archichlamydae 39 Betacyanins 123 Briar-pipes 368
Babool 302
Archichlamydeae 28 Betalains 121, 123 Bridelia 447
Baccharis 356
Arctostaphylos 366 Betaxanthins 123 Brinjal 400
Bacillariophyta 28
Arctotis 360 Betel 443 Broad bean 306
Bacopa 407, 551
Areca 500, 503 Betel-nut palm 503, 500 Bromelia 479, 480, 482
Badam 315
Arecaceae 52, 500, 504, 546 Betula 80 Bromeliaceae 479, 482, 545
Bael 279
Arecidae 39 Betulaceae 101 Bromeliaceae1 469
Baer 286
Arenaria 250 Bhang 459 Bromus 136
Bahera 323
Arenga 502, 503 Bhindi 260 Broom grass 523
Bahia grass 518
Argemone 124, 400, 528, 548 Bhrangraj 359 Broussonetia 454
Baingan 400
Argyreia 394 Bibliography 149 Brunfelsia 399
Bajra 518, 522
Arhar 309 Bicarpellatae 25 Brunoniaceae 355
Bakain 282, 429
Aril 292 Bidens 356 Brussel’s sprouts 245
Balanophoraceae 29, 72
Arisaema 507, 508, 509 Biennial 176 Bryonia 331
Balanophorales 29
Aristida 518, 524 Biflavonyls 122 Bryophyta 27
Balanophoreae 26
Aristolochia 9, 97 Bifoliate 190 Bsi 16
Balanopsidaceae 28, 429
Aristolochiaceae 29, 544 Biggest Indian herbarium 152 Buchanania 293
Balanopsidales 28
Aristolochiales 29 Bignonia 411 Bucida 322
Balausta 206
Aristotelian Concept 49 Bignoniaceae 25, 409, 544 Buckwheat 437
Baliospermum 447
Aristotle of the Middle Ages” 9 Bignoniaceae, Pedaliaceae 34 Bud 176
Balloon vine 290
Arjun 322 Bilabiate 169, 198 Buddleia 124
Bamboos 523
Armeria 372 Billbergia 479, 480, 481, 482 Buddleiaceae 124
Bambusa 518, 519, 520, 523, 524
Arrabidaea 409 Bilva 279 Buettneria 263
Banafsha 249
Arrowhead 510 Binary Characters 116 Bulb 179
Banana 486
Arrowroot 450 Binomial 48, 89 Bulbel 179
Band Gobhi 244
Artabotrys 218, 429 Binomial system of nomenclature 10, Bulbil 179
Bankla 309
Artemisia 356, 550 21 Bulbophyllum 470
Bank notes 453
Artificial Classifications 19 Biological Concept of Species 50 Bulbostylis 513
Banyan tree 455
Artificial Keys For The Identification Biophytum 274 Bull grass 518
Baphia 100
72 Biosystematics 1 Bupleurum 342
Baptisia 122
Artocarpus 454 Birch 80 Burmanniaceae 28, 469
Bara Nimbu 279
Arum 507, 508 Biscayne palm 500 Burseraceae 280
Barbacenia 100
Arundinaria 518, 523 Bischofia 450 Butea 310
Barbarea 245
Arundo 524 Bitter gourd 331 Butein 122
Barclaya 228
Arvi 509 Bixineae 231 Butolaceae 29
Barclayaceae 229
Asafoetida 343 Blackberry 316 Butomaceae 34
Bargad 455
Asarum 438 Black gram 309 Butomus 111, 510
Barhal 455
Ascending 179 Black pepper 443 Buttercup 166
Bari Champa 215
Asclepiadaceae 25, 29, 380, 534, 543 Blastophaga 455 Butter-cup 223
Barleria 414, 331
Asclepias 380 Blepharis 414 Button Gobhi 245
Barley 518, 520
Asepalous 196 Blighia 292 Buttress 177
Barosma 280, 548
Asgandh 401 Blumea 359, 550 Buxaceae 289
Barringtonia 324, 172
Ashok 219 Bocconia 232
Index 555
Cabbage 244 Carpel 65, 245 Chakotra 278 Cistaceae 106

Cabbage palm 500, 502, 503 Carpet grass 523 Chamaecrista 303 Cistineae 231
Cabomba 227, 29 Carrisa 386 Chamaedorea 500 Cistus 106
Cabombaceae 229, 32 Carrot 343 Chamaerops 502 Citation 149
Cacervulus 206 Carrot grass 359 Chamaesyce 100 Citation of Author 87
Cactaceae 25, 72, 542 Carthamus 359, 221 Champa 215, 377 Citronella oil 523
Cactales 32, 30 Carum 343 Chana 308 Citrullus 330, 436
Cactus 10 Caryophyllaceae 25, 37, 249, 540 Chandni 387 Citrus 123
Cadaba 240 Caryophyllales 31, 350, 174, 98 Characteristics of the angiosperms 138 Clades 91
Caducous 196 Caryophyllidae 39, 255 Characters 46 Cladistics 130, 518
Caesalpineae 542 Caryophyllineae 25, 213, 460, 461, 462 Character states 46 Cladium 517
Caesalpinia 303, 222, 213, 549 Caryopsis 206, 520 Charaka Samhita, 13 Cladogram 130
Caesalpiniaceae 173, 531 Caryopteris 425 Charas 459 Cladophyll 179
Caesalpinioideae 74, 255, 299, 302 Caryota 500, 309 Charophyta 28 Clarkia 105
Cajanus 307, 167 Cashew-nut 293, 296 Chartacalyx 266 Clasping or Climbing 177
Cakile 125 Cassava 449, 450 Chasmanthium 524 Class 52
Caladium 509 Cassia 303, 549 Chaulai 434 Classification 4, 19
Calamus 500, 503 Castilla 455 Chawal 521 Claytonia 254
Calandrinia 254, 195 Castilleja 405, 124 Cheiranthus 244, 331 Clematis 70
Calceolaria 405, 324 Castor bean 446 Chelidonium 232 Cleome 86, 548
Calendula 359, 27 Castor oil 449 Chelone 406 Clermontia 363
Californian Poppy 233, 396 Casuarina 99, 136, 113 Chemical Plant Taxonomy 121 Clerodendrum 423
Callicarpa 423 Casuarinaceae 28, 282, 99, 460, 545 Chemical Taxonomy 111, 121 Clethraceae 365
Callistemon 323, 40 Casuarinaceae1 429 Chemosystematics 121 Clianthus 309
Calocarpum 377 Catalpa 409, 374 Chemotaxonomy 111, 122, 266 Climber 176
Calonyction 395, 362 Catechu of Commerce 302 Chenopodiaceae 25, 544 Climbing Ylang-Ylang 219, 220
Calotropis 380, 174, 527, 550 Categories of systems of classification Chenopodiaceae, Amaranthaceae 34 Clitoria 307
Caltha 221, 124, 333 19 Chenopodiales 253 Clivia 478
Calycanthaceae 102, 32, 396 Catharanthus 168, 550 Chenopodium 119, 551 Clove 326
Calyceraceae 355, 219 Catkin 192 Cherry 312 Clover 306, 309
Calyciferae 34 Cattleya 470 Cherry laurel 316 Cluster analysis 118
Calyciflorae 25, 110, 75 Caudate 182 Chewing gum 376 Cluster bean 309
Calycineae 26, 469 Caudicle 472 Chiananthus 380 Clutia 446
Calycopteris 323 Caulescent 179 Chichenda 331 Cneoraceae 280
Calystegia 97, 348, 241 Cauliflower 244, 245 Chicle 376 Cobaea 402, 331
Calyx 64, 195 Cauline 184 Chicle gum 377 Coccinia 330
Campanales 25, 53, 362 Cautleya 488, 103 Chikrassia 282 Coccoloba 435
Campanula 166, 241 Cayratia 286, 240, 549 Chiku 377 Coccothrinax 500
Campanulaceae 25, 411, 52, 543 Caytonialean Theory 142 Chillies 400 Coconut 503
Campanulales 32 Ceanothus 285, 400 China rose 260 Coconut palm 500, 502
Campanulatae 30 Cecropia 451 Chinese water-chestnut 516 Cocos 500
Campis 409, 29 Cedrela 282 Chionanthus 378 Codex Juliana. 9
Cananga 218, 240 Ceiba 269 Chirchita 435 Codiaeum 447
Canarina 363 Celastraceae 286 Chironji 296 Coding of characters 117
Canary grass 523 Celastrales 25 Chlaenaceae 257 Codonopsis 363
Candollea 80 Celery 343 Chlidanthus 478 Coefficients of association 117
Cane 503 Celery-leaved Crow-foot 224 Chloranthaceae 101 Coefficients of correlation 117
Cane or Rattan palms 503 Cellulares 23 Chlorophyceae 28 Coffea 349, 350
Cane palm 500 Celosia 432 Chlorophytum 97, 305 Cohorts 27, 366
Canna 482 Cenchrus 523 Chloroxylon 280 Coix 523
Cannabinaceae 174, 22, 545 Centaurea 356 Choice of Names 88 Cola 263
Cannabis 457, 251, 482 Centella 103, 550 Choisya 280 Colchicine 493
Cannaceae 108, 111, 400, 545 Centrantherum 359 Choriophyllum 446 Colchicum 490, 551
Cannellaceae 231 Centrolepidaceae 470 Chorizanthe 435 Coleus 419
Canon of Medicine, 9 Centropogon 362 Chosenia 463 Collecting Bags 56
Capers 241 Centrospermae 29 Choti Elayachi 489 Collection Bottles 56
Capitulum 356 Cephaelis 349, 550 Christmas rose 223 Collector number 56
Capparaceae 125, 249 Cephalanthus 350 Chromosome number 103 Colletia 285
Capparales 241, 400 Cephalocereus 339, 523 Chronology 14 Collinsia 407
Capparidaceae 24, 30, 74, 239, 540 Cephalostachyum 523 Chrozophora 447 Collomia 402, 508
Capparis 240, 89 Cerastium 250 Chrysanthemum 355 Colocasia 507
Caprifoliaceae 25, 28, 543 Ceratonia 303 Chrysobalanus 314 Columelliaceae 405
Capsella 98, 243, 216 Ceratophyllaceae 26, 223, 429 Chrysophyllum 375, 270 Column 472
Capsicum 399, 304, 360 Ceratostigma 372 Chrysoplenium 318 Combretaceae 25, 321, 542
Capsule 206 Ceratotheca 413 Chufas 516 Combretum 321, 495, 496, 549
Caralluma 382 Cerbera 385, 286 Chukander 429 Commelina 494, 381, 382, 497
Carameriana 287 Cercidiphyllaceae 213 Chukrasia 284 Commelinaceae 26, 28, 546
Carapa 282 Cercis 303 Cicer 108, 424, 425 Commelinaceae1 469
Caraway 343 Cercocarpus 80 Cichorium 357 Commelinidae 39
Cardamine 243 Cercopia 452 Cicuta 343 Common names 79
Cardamon 489, 490 Cereus 339 Cimicifuga 136, 345 Common Prefixes 89
Cardiospermum 290 Ceropegia 381, 237 Cinchona 162, 343, 345, 344, 550 Common Suffixes 89
Carduus 356 Ceroxylon 503 Cincinnus 192, 392 Comparison of systems of classifica-
Carex 100 Cespitose 179 Cirrhose 183 tion 36
Carissa 387 Cestrum 399, 309 Cirsium 356 Complete 195
Carnation 250 Cgiar 59 Cissus 286 Compositae 12, 543
556 Index
Compound 192 Crinum 477 Cyrtanthus 477 Dictamnus 277, 549

Compound Leaf 189 Crocosmia 476 Cytinaceae 429 Didieraceae 123
Computerised identification 71 Crocus 474 Cytology 103 Didymous 201
Concept of Ecological Species 50 Crossandra 414 Cytotaxonomy 103 Didynamous 201
Concept of Evolutionary Species 50 Crotalaria 119, 549 Czekanowskiales Theory 143 Dieffenbachia 509
Concept of family 52 Croton 446 Die Naturlichen Pflanzenfamilien 11
Dactylis 523
Concept of Genus 51 Crown jewels 59 Diervilla 354
Dactylorhiza 474
Concept of “minimum invalidity 85 Cruciales 245 Differences between dicots and mono-
Daemonorops 500, 503
Concept of selection species 51 Cruciferae 22, 243 cots 139
Daffodil 479
Concept of Sibling Species 50 Cruciform 199, 243 Diffuse 181
Dahlia 355, 399, 400
Concept of Species 47 Cryptangium 518 Digera 432, 537
Dalbergia 97, 71
Concept of Taxa 46 Cryptantha 392 Digitalis 99, 551
Damasonium 510
Condalia 285 Cryptanthus 482 Digitaria 518
Danthonia 524
Conical 177 Cryptogamae 27 Dill 343
Daphnales 26
Coniferae 26 Cryptogamia 21 Dilleniaceae 213
Darwinia 324
Coniferales-Amentiferae Theory 142 Cryptogams) 8 Dilleniidae 37
Date palm 500, 503
Conium 121 Cryptostegia 380 Dimorphotheca 359
Datura 136, 221, 222, 550
Conjugatae 28 Cubebs 443 Dinoflagellatae 28
Daucas, 8
Connaraceae 100 Cucubalus 251 Dionysia 369
Daucus 342, 304
Connective 200 Cucumber 330 Dioscorea 97
Debregeasia 453
Contortae 29 Cucumis 330 Dioscoreaceae 102, 469, 482
De Candolle rules 81
Contractile 177 Cucurbita 119 Diplanthera 409
Decliptera 414
Convolvulaceae 25, 543 Cucurbitaceae 25, 329, 542 Diploknema 377
Decompound 191
Convolvulus 394 Cucurbitaceae, Begoniaceae 34 Diplostemonous 201, 303
Decumbent 181
Copaifera 304, 503 Cucurbitales 30 Dipsacaceae 124
Deep-freeze Arctic Vault 59
Copernicia 500 Cullenia 269 Dipsacales 44
Deep-freezing methods 58
Coperonia 446 Culm 179, 514 Dirachma 273
Deeringia 432, 320, 536
Coprosoma 349 Culms 519 Discaria 286
Definable families 52
Coptis 105, 449, 547 Cultivar 48 Disc florets 356
Degeneriaceae 101
Corallorhiza 471 Cultivated variety 48 Dischidia 381
Deinbollia 290
Corchorus 265, 365, 548 Cumin 343 Disciflorae 25
Delonix 303, 72
Cordia 392 Cuminum 343 Distichia 497
Delphinium 70, 99, 106, 527, 547
Coreopsis 359 Cuphea 327, 479 Distichous 186
Dendrobium 470, 471, 474
Coreya 326 Curculigo 478, 489, 551 Division 19, 52
Dendrocalamus 519, 523
Coriander 343 Curcuma 488, 489, 551 Dizygotheca 348
Dendrogram 130
Coriandrum 342 Curvembrae 27 DNA barcoding 128
Dendromecon 232, 345
Corispermum 430 Curvembreae 428 DNA-DNA hybridisation 129
Dendrophthoe 443
Corm 179 Curvembryeae 25 DNA-fingerprinting 132
Dendrophthora 443
Cormel 179 Cuscuta 394 DNA-markers 132
Dendrophylax 472
Cornaceae 85 Cuscutaceae 397 DNA-sequencing 129
Dendrosicyos 330
Cornales 44 Cusparia 277, 549 Dobinea 293
Derris 310
Corolla 64 Cuspidate 183 Doctor Universalis” 9
Description 5
Corolliferae 34 Cuticular papillae 98 Document 148
Desfontaineaceae 377
Corona 336, 469 Cyamopsis 309 Documentation 148
Desmanthus 299
Coronarieae 26 Cyananthus 363 Document Retrieval 149
Desmodium 306
Coronopus 243, 472 Cyanea 362 Docynia 317
Desmoncus 500
Corpusculum 381 Cyanide 338 Dodder 395
Deutzia 318, 300
Correa 277 Cyanobacteria 81 Dodecatheon 369
Dewberry 316
Corrugate Ventilators 56 Cyanogenesis 126 Dodonaea 289
Dewey Decimal (DD) system 149
Cortaderia 523 Cyanogenic compounds 126 Dolichandrone 409
Dhaincha 310
Corydalis 236, 548 Cyanophoric plants 126 Dolichos 306, 424, 425
Dhak 310
Corylaceae 443 Cyanotis 494 Dombeya 265
Dhania 343
Corymb 192 Cyathium 192, 446 Doob grass 518, 523
Dhobis-nut 296
Corypha 503 Cyathocalyx 218 Dorstenia 454
Diadelphous 200
Cosmos 359 Cycadaceae 26 Douglasia 371
Dialium 303
Costaceae 108 Cyclamen 369, 470 Draba 243
Dianella 491
Costus 488 Cyclanthaceae 28 Dracaena 490, 493
Dianthus 250, 548
Cotinus 293 Cyclanthera 331 Dracontium 507
Diapensiaceae 29
Cottea 524 Cydonia 317 Dracunculus 509
Diapensiales 29
Cotton 260 Cylindrical 181 Drier 56, 58
Diarrhena 524
Cotyledon 22, 184 Cymbidium 470 Drimycarpus 296
Dicentra 236
Co-types 87 Cymbopogon 523 Drimys 144
Dichanthium 523
Couplet 70 Cymose 192 Droseraceae 108
Dichasium 192
Cousinia 356 Cynara 359 Drupe 206
Dichondra 397
Cowpea 309 Cynoctonum 389 Drying chamber 58
Dichondraceae 397
Crab grass 518 Cynodon 518 Drying of Specimens 57
Dichopsis 375
Crambe 244 Cynoglossum 392 Drymaria 97
Dichorisandra 494, 495
Cranberry 355 Cynosurus 523 Drymis 145
Dichotomous 181
Crassulaceae 110 Cyperaceae 26, 513, 514, 515, 516, 517, Drypetes 450
Dichotomous keys 70
Crataegus 312 518, 546 Duboscia 266
Diclidantheraceae 375
Crataeva 240 Cyperus 98, 551 Dulee Champa 215
Dicotyledoneae 13
Creeper 176 Cyphia 362, 471 Dulichium 518
Dicotyledones 20, 434, 435
Cremocarp 343 Cypripedium 470 Duranta 423
Dicotyledons 12, 24, 74
Cremocarp 206 Cypsela 355 Durian Theory 143
Dicrostachys 299
Crepis 356 Cypsela 206 Durio 143, 548
Dicta 12, 30
Crescentia 409, 478 Cyrtandromoea 103 Dyckia 480
Index 557
Dysoxylum 282, 549 Ericales 25 Fibrous 177 Gamotepalous 199

Dysphania 250 Erigeron 356 Ficoidales 25 Ganja 459
Erinocarpus 267 Ficus 454 Ganna 522
Ebenaceae 375, 377 Eriobotrya 312 Field equipment 55 Ganth Gobhi 244
Ebenales 25, 29, 374 Eriocaulaceae 470 Field notebook 56 Gardenia 349
Ecballium 331 Erioglossum 290 Field preparation of specimens 55 Garden pea 308
Echinocactus 339, 340 Eriogonum 435, 536 Field Press 56 Garlic 493
Echinocereus 339 Eriolaena 265 Fig 455 Garryaceae 28
Echinochloa 519 Eriostemon 277 Figwort 405 Garryales 28
Echinocystis 330 Eritrichium 392 Fimbristylis 104, 513, 514 Gart der Gesundheit” 9
Echinodorus 510, 513 Erodium 271 Firmiana 265 Gasteria 491
Echinops 356 Eruca 243 Fish-tail palm 500, 502 Gaultheria 366, 27
Echinopsis 339, 360 Erycibe 395 Fittonia 415 Gaylussacia 366
Echium 393 Eryngium 342 Flacourtiaceae 125 Gehun 521
Eclipta 355, 29 Erysimum 243 Flagellariaceae 26, 469 Gelonium 450
Edrianthus 363 Erythrina 306 Flagellatae 28 Gelsemium 391
Effectively published 88 Escallobonia 318 Flavones 122 Genda 359
Effecttve and Valid Publication 88 Escallonia 320 Flavonoids 122 Genealogy 138
Eggfruit 377 Eschscholzia 231 Flavonols 122 Gene Bank search 131
Eggplant 400 Essence 49 Flavonones 122 Genera Plantarum 21
Ehretia 392 Eucalyptus 97 Fleshy 177 Genera Plantarum. 11
Eichhornia 101 Eucharis 478 Flindersia 277 Generic epithet 79
Eichleria 274 Eucryphia 123 Flora 148 Generic Name 80
Elaeagnaceae 429 Eucryphiaceae 123 Floral anatomy 102 Genetic fingerprinting 129
Elaeis 503 Eugenia 324, 549 Floral characters 97 Geniculate 181
Elaeocarpaceae 257 Eugenia complex 51 Floral diagram 67, 343 Geniostoma 389
Elaeocarpus 265 Eulophia 474 Floral formula 67, 237 Genista 125
Elaiosome 247 Eumycetes 28 Floral Symbols 166 Genome 129
Elaisome 222 Eupatorium 356 Florets 519 Gentianaceae 106
Elatostema 451 Euphorbia 22 Floridean royal palm 502 Gentianales 25, 405, 280
Elayachi 489 Euphorbiaceae 26, 537, 545 Floscopa 495 Genus 19, 279
Eleocharis 513, 514 Euphorbiaceae 445 Flower 64 Genus; 19
Elephant foot 509 Euphorbiaceae1 429 Flower 176 Geraniaceae 25, 308, 309, 529, 541
Elettaria 488, 489 Euphorbiales 43 Flowering plants 8, 138 Geraniales 25, 402, 404
Eleusine 522 Euphoria 292 Foeniculum 342 Geranium 271, 169, 548
Eleuthrine 475 Euphrasia 406 Follicle 206 Geranium oil 273
Emblica 446, 341, 551 Euploidy 104 Forget-me-not 394 German Fathers of Botany” 9
Emblingia 240, 245 Euptelea 215 Form 48 Gesneriaceae 103
Embryology in solving taxonomic prob- Eupteleaceae 216 Formalin-aceticacid-alcohol (FAA) 56 Geum 314, 532
lems 109 European fan palm 502 Forskohlea 452 Geunsia 424
Embryophyta 28 Euryale 227 Forsythia 380 Gherkin 331
Embryophyta-Siphonogama 28 Eurypalynous 107 Fortunella 279 Ghia Tori 331, 309
Empetraceae 429 Evodea 277 Fossil angiosperms 140 Ghikanwar 493
Empetrum 22 Evodia 280 Fountain palm 502 Ghuiyan 509
Endings of ranks 5 Evolutionary trees 130 Foxglove 405 Gigantochloa 523
Engler and Prantl’s system 27 Evolvulus 394 Foxtail 434 Gigardinia 453
Ensete 485, 486, 487 Examination of a plant specimen 61 Foxtail millet 523 Gilia 401
Entada 299, 228 Exocarpaceae 111 Fragaria 312, 532 Gingelly 412
Epacridaceae 98 Exocarpus 111 Francoa 318 Ginger 489
Ephedra 460 Exogonium 396, 550 Frangula 285 Ginger grass 523
Epicalyx 184, 258, 367 Exstipulate 184 Frankenia 101 Glabrous 181
Epidendrum 471, 474 Extrorse 169, 201 Frankeniaceae 249 Gladiolus 474, 475
Epigaea 366, 323 Fraxinus 378 Glaucous 181
Fabaceae 52, 74, 124, 173, 297, 307
Epigynae 26, 469 Free-central 205 Glaux 371
Fabales 297
Epigynous 22 Freesia 475 Gleditsia 303
Fagaceae 29
Epigynous 195, 325 Fritillaria 490, 493 Glinus 97
Fagales 29
Epipendrum 470 Fruit 67, 176 Globba 488, 489, 490
Fagara 277
Epipetalous 169 Fuchsia 123 Glochidia 339
Fagopyrum 22, 435, 437
Epipetalous 201 Fuirena 100 Gloriosa 491, 493, 551
Fagraea 389
Epiphyllum 339 Fumaria 236, 528, 548 Glossopteridalean 142
Family 19
Epiphyte 177 Fumariaceae 109, 236, 528, 540 Glossopteris 97
Family tree 136
Epipogon 472 Fungi 22, 27 Glucosinolates 125
Farinosae 28
Epithets Linked with Colour 90 Furcraea 477, 479 Glumaceae 26, 470
Fasciculated 177
Epithets Linked with Geography 90 Fusiform 178, 243 Glume 184
Father of Indian Botany 162
Epithets Linked with Habit 90 Future of Plant Taxonomy 5 Glumes 519
Father of taxonomy 21
Epithets Linked with Habitats 91 Glumiflorae 28
Fatsia 348 Gaertnera 350
Epithets Linked with Size 90 Glycine 306
Feather grass 523 Gahnia 514
Equisetum 460 Glycosmis 277
Feijoa 325 Gaillardia 359
Eragrostis 518, 524 Glycyrrhiza 549
Fennel 343 Gajar 343
Eranthemum 414 Gmelina 425
Fernary 162 Galanthus 477, 478
Eranthis 222 Gnaphalium 356
Ferns 22 Galgal 278
Erect 181 Gnetaceae 26
Feronia 277, 549 Galium 137, 32
Eremostachys 100 Gnetales-Angiosperm Theory 143
Ferula 342, 549 Gamopetalae 25, 51
Erianthus 523 Goat weed 361
Festuca 100, 523 Gamopetalous 197
Erica 366, 359 Godetia 105
Fevillea 330 Gamosepalous 196
Ericaceae 25, 543 Gomphrena 432
558 Index
Gomuti palm 502 Hastate 187 Huckleberry 366 Iridaceae 26, 469, 477, 482, 538, 545
Goniothalamus 219 Haustorial 178 Huernia 382 Iridaceae 474
Gonophyll theory 102 Haworthia 490, 491 Humulus 457 Iridales 31
Goodeniaceae 355 Head 356 Hura 450 Iridoids 124
Goolar 455 Head or Capitulum 192 Hurhur 241 Iris 21, 474, 475, 476, 551
Gossypium 257, 548 Heath 367 Hybanthus 247, 548 Isatis 244, 350
Gouania 285 Hedera 347, 393 Hydrales 31 Iseilema 523
Gram 308 Hedychium 488, 489, 490 Hydrangea 318 Isoetes 141
Graminae 76 Heeng 343 Hydrastis 136 Isoetes-Monocotyledon Theory 141
Graminales 31 Helianthemum 106 Hydrocaryaceae 111 Isoflavones 122
Gramineae 26 Helianthus 355 Hydrocharitaceae 469 Isoflavonoids 122
Gramineae1 470 Helichrysum 356 Hydrocotyle 103 Isonym 84
Grandfather of the modern botany” 8 Heliconia 485, 486, 487 Hydrophyllaceae 391 Isophysis 475
Grape vine 287 Heliconiaceae 108, 486 Hydrostachyaceae 28 Isopyrum 105
Grayia 430 Helicteres 262, 400, 548 Hydrostachyales 28 Isotype 86
Great Banyan tree 162 Helinus 285 Hygrophila 415, 551 ISPN 93
Green gram 309 Heliophila 243 Hymenaea 304 Italian millet 518, 522
Grewia 265, 548 Heliotropium 392 Hymenocallis 477, 479 Ivory-nut palm 500
Grindelia 359, 550 Helleboraceae 224 Hyobanche 406 Ixora 349
Groundnut 308 Helleborus 221, 547 Hyoscyamus 136
Jacaranda 409, 411
Guarana 292 Helobiae 28, 237 Hypanthium 195
Jack fruit 455
Guava 325 Helwingia 346 Hypanthodium 192, 194
Jacobinia 415
Guazuma 264 Hemerocallis 491, 493 Hypecoum 236, 548
Jacquemontia 395
Guettarda 349 Hemicyclia 450 Hypericaceae 52
Jaculator 415
Guide to the examination of plant speci- Hemidesmus 384 Hypericum 22
Jai 520
men 62 Hemp 260, 459 Hyphaene 503
Jalap 396
Gulkand 317 Henbane 400 Hypochoeris 356
Jamalghota 450
Gulmohar 304 Henna plant 329 Hypocrateriform 199
Jambolan 325
Gum arabic 302 Heracleum 345 Hypogynous 22, 195
Jamun 325
Gutta percha 377 Herb 177 Hypoxis 477, 479
Jangli Khajoor 503
Guttiferae 25 Herbaceae 34 Hyptis 419
Jangli Posth 233
Guttiferales 25 Herbae 20
Iberis 243, 244, 245 Jasione 365
Gwar 309 Herbalists 9
Icacinaceae 99 Jasmine 350, 447
Gymnema 383 Herbals 9
ICBN 78 Jasminum 378
Gymnocladus 305 Herbarium 152
ICNCP 81 Jatropha 446, 551
Gymnospermae 26 Herbs 20
Idenburgia 101 Jaun 520
Gymnosperms 72, 138 Heritiera 264
Identification 4, 69 Jeans 459
Gynandrium 381 Hermaphrodite 166, 194
Identification Methods 72 Jerusalem artichoke 359
Gynandropsis 86 Hermodactylus 475
Identification of plants 60 Jhar 286
Gynandrous 201 Herpestis 407
Identification With Keys 70 Job’s tears 523
Gynoecium 65 Hesperelaea 378
Ideographs 166 Joint Author 87
Gynophore 240 Hesperidium 206
Idiogram 103 Jowar 522
Gynophore 195 Heterocontae 28
Illeceraceae 428 Juglandaceae 29
Gynostegium 381 Heteromerae 25
Illiciaceae 137 Juglandales 29
Gynostemium 440 Heterophragma 409
Illicium 137, 78, 80, 547 Julianiaceae 29
Gynura 359 Heteropogon 523
Imbricate Aestivation 196 Julianiales 29
Gypsophila 250 Heterotypic synonym 84
Imparipinnate 190 Juncaceae 26, 469, 497, 498, 517, 538,
Heuchera 320
Habenaria 470, 473 Imperfectae 20 539, 546
Hevea 446
Habit 177 In 97 Juncaginaceae 126
Hibiscus 257, 548
Habitat 177 Inadmissible characters 116 Juncus 99, 497, 498, 499
Hieracium 356
Haemanthus 477, 478, 479 Incarvillea 411 Justicia 414
Hildebrandtia 395
Haematoxylin 304 Indefinable families 52 Jute 265
Hillebrandia 333
Haematoxylon 304 Indented or yoked key 70
Himantandraceae 214 Kaans 523
Haemodoraceae 469, 482 Indian botanical libraries 150
Hina 260 Kababchini 443
Haemodorum 492 Indian Journals 13
Hippeastrum 477, 478, 479 Kachalu 509
Hagenia 317, 549 Indian rape 245
Hippocastanaceae 297 Kachnar 304
Haldi 489 Indian rubber plant 455
Hirtella 312 Kachra 331
Haldu 352 Indigo 310, 306
History and Development of Plant Kadam 350
Haloxylon 430 Indigofera 306
Classification 8 Kaempferia 488, 490
Hamamelidaceae 297 Indigo plant 309
History of ICBN 81 Kaghzi Nimbu 278
Hamamelidae 39, 264 Inferae 25
Hog plum 293 Kaith 279
Hamamelididae 37 Inferior Ovary 202
Holarrhena 385 Kakdi 331
Hamelia 349, 177 Inflorescence 64
Holigarna 293 Kala Shisham 309
Hamiltonia 350 International Botanical Congresses 78
Holmskioldia 425 Kala Zeera 224
Hancornia 387 Inter national Code of Botanical
Holotype 86 Kali Rai 245
Hanging Gardens 158 Nomenclature 4, 360
Holy basil 421 Kali Tori 331
Haplopappus 103 International Code of Nomenclature of
Homonym 87 Kalmia 367
Haplophragma 411 Bacteria (ICNB) 82
Homoploids 104 Kalonji 224
Haplotype 130 Introrse 169
Homotypic synonym 84 Kalopanax 348
Haptens 135 Inula 356, 395, 550
Hoplophyllum 356 Kamal 228
Hararh 323 Ipecac 350
Hops 459 Kamal-Kakri 229
Hardwickia 304 Iphigenia 493
Hordeum 126, 518, 520 Kanduri 331
Hare’s-tail grass 523 Ipomoea 394, 434, 435
Hortus Sanitalis” 9 Kaner 387
Harpephyllum 296 Iresine 432
Hovenia 286 Kantali Champa 220
Hashish 459 Iriartea 500
Hoya 381 Kapok 269
Index 559
Karela 331 Lavender 419 Lipids 125 Major Botanical Gardens of India 161
Karonda 387 Lawn Grasses 523 Lippia 423 Major Botanical Gardens of The World
Karyotype 103 Lawsonia 327 Liquid Preservative 56 158
Katha 302 Lead 70 Liriodendron 214 Major Botanical Libraries 150
Kathal 455 Leaf 63 Lissocarpaceae 375 Major herbaria of the world 154
Keel 307 Leaflet 184 Litchi 289 Major Indian herbaria 154
Kela 486 Leaf shapes 188 Literature Retrieval 148 Makhana 229
Keys 69 Leaf venation 191 Lithospermum 392, 550 Makka 522
Kharbooja 331 Lectotype 86 Littorella 426 Makoi 401
Khatta 278 Leea 286, 215, 549 Living Fossils 144 Malachra 260
Khaya 282 Legume 209 Livistona 501, 502 Mallotus 450, 75
Kheera 331 Leguminales 297 Loasales 32 Malta 279
Khirni 377 Leguminosae 25 Lobelia 362, 550 Malus 123, 173
Khubani 316 Lehsua 435 Lobia 309 Malva 257, 29, 32, 548
Khus-Khus 523 Leitneriaceae 29 Lobularia 245 Malvaceae 22, 214, 51, 529, 541
Kigelia 409 Leitneriales 29 Lodicule 197, 519 Malvales 25, 75, 97, 101
Kikar 302 Lemma 519 Logania 389 Malvastrum 257, 32
Kingdom 52 Lemnaceae 28, 470, 504, 509 Loganiaceae 124, 543 Malvaviscus 260
Kirganelia 450 Lemon 278 Loganiales 391 Mammillaria 339, 281
Kmeria 214 Lens 309 Lolium 523 Manadenium 446
Knol-knol 245 Lentibulariaceae 405 Lomentum 209 Mandevilla 387
Kochia 429 Lenticel 181 Long pepper 443 Mandragora 400, 550
Koenigia 436 Lentil 309 Lonicera 352 Mangifera 293
Kolkwitzia 354 Lepidium 243 Loofah 330 Manglietia 214, 449
Korthalsia 503 Leptadenia 380 Lophophora 341 Mangnoliaceae 213, 414
Krameria 109, 549 Leptopyrum 136 Loquat 317 Mango 293, 296
Krameriaceae 109 Leptospermum 326 Loranthaceae 26, 443, 544 Manihot 446, 453, 329
Kusum 359 Lesquerella 106 Loranthaceae1 429 Manila palm 502
Kuttu 437 Leucaena 299, 251 Loranthaceae, Santalaceae 34 Manilkara 375, 102
Kydia 258 Leucas 419 Loranthoideae 110 Manioc 450
Kyllinga 515 Leucocarpus 406 Loranthus 443 Mankanda 509
Leucojum 479 Lotus of India 228 Mansonia 263
Label 59
Leucomeris 356 Love grass 518 Maoutia 451
Labelling of Specimens 59
Leucosceptrium 419 Love-in-a-mist 223 Mapania 518
Labellum 472
Leucothoe 367 Lowiaceae 486 Marantaceae 483
Labiatae 25, 112, 121, 173, 419, 535,
Lewisia 254, 377 Lridaceae 174 Marginal 205
544
Leycesteria 354 Luffa 330 Marigold 359
Lablab 309
Liabum 360 Lumnitzera 321 Marijuana 459
Lacistemaceae 429, 356
Liana 177 Lunaria 244 Mariscus 515
Lacquer tree 296
Library of Congress (LC) system 149 Lupine 306 Marrubium 101
Lactuca 356, 328, 30
Lichens 27 Lupinus 124 Marsdenia 383
Lady’s finger 260
Lightfootia 362 Luvunga 280 Marsippospermum 497
Lagenaria 330, 126, 39
Lignosae 34 Luzula 497, 498 Martynia 412
Lagerstroemia 327, 296
Ligularia 356 Lychnis 250 Martyniaceae 414
Lagurus 523
Ligule 184, 519 Lycium 399 Masoor 309
Lahsun 493
Ligustrum 378 Lycopersicon 399 Matar 309
Laila Majnu plant 326
Lilac 380 Lycopodium 124 Matelea 381
Lamarckia 523
Lilaea 103 Lyonia 366 Materia Medica 9
Lamiaceae 109, 38, 544
Lilaeaceae 103 Lysichiton 509 Materia Medica. 8
Lamiaceae 419
Liliaceae 26, 406, 477, 479, 490, 493, Lysimachia 369 Mathiola 245
Lamiales 25, 258
494, 498, 538, 546 Lythraceae 25, 542 Maulsari 377
Laminar 205
Liliaceae1 469 Lythraceae 327 Maurandia 406
Lamium 419
Liliales 31 Lythrum 327 Mayaceae 469
Landolphia 385, 32
Liliatae 469 Mazus 405, 255
Lannea 293, 29 Mackinlaya 346
Liliidae 13, 331, 332, 333, 469 Mechanical Classifications 20
Lantana 423 Macleaya 232, 241
Liliiflorae 28 Meconopsis 231, 241
Laportea 101 Maclura 454, 455
Liliopsida 12, 469 Medicago 306
Lappula 392 Macropiper 443
Lilium 490, 491, 493 Meetha Nimbu 278
Lardizabalaceae 213 Madar 384
Lily 493 Megacarpaea 244
Largest Botanical Garden of India 162 Madder 352
Lima bean 309 Mehndi 329
Largest Botanical Garden of The World Madhuca 375, 39, 29, 550
Lime 278 Melaleuca 324, 331
163 Maerua 240, 13
Limnophyton 510 Melastomaceae 101, 331
Larkspur 223 Magnolia 136, 25, 43
Limonia 276 Melia 281, 549
Lasia 507 Magnoliaceae 24, 29, 443, 513, 526,
Limoniaceae 100 Meliaceae 25, 281, 530, 541
Lassora 394 540
Limonium 100, 394 Melica 524
Lathrea 406 Magnoliaceae 214
Linaceae 111, 328 Melicocca 292
Lathyrus 125 Magnoliales 144
Linanthus 404 Melilotus 307
Laticiferous 181 Magnoliatae 212
Linaria 405 Melocanna 523
Latin Diagnosis 88 Magnoliidae 13
Lindenbergia 405 Melochia 262
Latjeera 435 Magnoliiflorae 41
Lines of Evolution in Angiosperms 145 Melon 331
Lauki 331 Magnoliophyta 12, 37
Linnaea 10, 22, 428 Mendoncia 414
Launaea 355 Magnoliopsida 12, 28
Linnaeus 21 Menispermaceae 103
Laung 326 Mahua 377
Linnaeus of India 14 Mentha 21, 551
Lauraceae 102 Maianthemum 491, 492
Linnaeus’s system of classification 21 Mercurialis 447
Lavandula 419 Maize 518, 522
Linociera 378 Mertensia 392
Lavatera 257 Majorana 421
Lip 472 Metachlamydae 39
560 Index
Metachlamydeae 43 Mounting 58 Nepetia 97 Opposite superposed 186

Metroxylon 500, 503 Mucronate 183 Nephelium 292 Opuntia 339
Michelia 136, 547 Muehlenbeckia 435 Neptunia 299 Opuntiales 29
Microembryeae 25 Muhlenbergia 518 Nerine 479 Orange 279
Microrhamnus 285 Multicostate 191 Nerium 384 Orchidaceae 22, 469, 470, 474, 545
Microscope. 9 Multifoliate 190 Nicandra 400 Orchidales 31, 271
Microspermae 26, 469 Multiovulatae-aquaticae 25, 429 Nicotiana 136, 550 Orchidantha 485, 486
Mikania 356 Multiovulate-terrestris 25, 429 Nidularium 482 Orchis 470, 473
Milkweed 381 Multipalynous 107 Niebuhria 240, 228 Orcuttia 524
Millet 522 Multistate Characters 116 Nigella 221, 547 Order 19
Milletia 310 Mung 309 Night Jasmine 400 Order 19
Millets 522 Munj 523 Nira 503 Order 52, 275
Millettia 307 Muntingia 268 Node 181 Ordines anomali 26
Millingtonia 409 Murdannia 494 Nodulose 179 Organisation of the field press 57
Miltonia 474 Murraya 276, 486, 487 Nolanaceae 397 Origanum 421
Mimosa 299 Musa 485 Nomenclatural Type 87 Original Author 87
Mimosaceae 108 Musaceae 28, 546 Nomenclature 5 Origin of Monocots 145
Mimoseae 541 Musaceae, Zingiberaceae 34 Nomenclaturist 78 Ornithogalum 490
Mimosoideae 74 Mussaenda 349 Nominalistic Concept of Species 48 Orobanchaceae 124
Mimulus 405 Mussambi 279 Nonflowering plants 8 Oroxylum 409
Mimusops 375, 550 Mustard 245 Nopalea 341 Orthosiphon 419
Mirch 400 Mutisia 357 Notalaea 380 Oryza 119, 518, 521, 523, 524
Mischodon 446 Mycorrhizal 178 Nothosaerva 433 Osmanthus 378, 437
Mitragyna 352 Myoporaceae 417 Nouhuysia 101 Out group 130
Mitreola 389 Myosotis 392 NPC-system 107 Ovary 202
Modern herbarium 153 Myosurus 136 Nucleic Acids 126 Ovule 203
Modern Trends in Plant Taxonomy 96 Myrcia 324 Nudiflorae 26, 470 Oxalidaceae 172, 274, 529, 541
Moghania 309, 549 Myricaceae 28 Number of carpels in a pistil 66 Oxalis 274
Molecular markers 131, 132 Myricales 28 Number-position-character analysis 107 Oxera 423
Molecular phylogenetics 128 Myriciaria 325 Numerical taxonomy 115 Oxychloe 497, 498
Molecular systematics 128 Myristicaceae 429 Nuphar 227 Oxymitra 218
Molecular taxonomy 128 Myristicta 101 Nut 209 Oxypetalum 381
Molecule 128 Myroxylum 310 Nuttalia 312 Oxyria 436
Molluginaceae 123 Myrsinaceae 369 Nuxia 389
Paan 443, 503
Mollugo 97 Myrtaceae 25, 542 Nuytsia 444
Pachycauly 181
Momordica 330 Myrtales 25 Nyctaginaceae 29
Pachylarnax 215, 216
Monadelphous 201 Myrtiflorae 29 Nyctanthes 99, 550
Paddy 521
Monandrous 201 Myrtus 323 Nymphaea 227, 547
Paederia 350
Monarda 421 Myxopyrum 378 Nymphaeaceae 24, 526, 540
Paeonia 103, 28, 547
Monera 47 Myxothallophyta 28 Nymphaeaceae 227
Paeoniaceae 103, 29, 73
Monerma 524 Nymphaeales 229
Najadaceae 470 Palaeobotany 112, 377
Money plant 507, 509 Nypa 500, 502, 503
Najas 142 Palak 429, 430
Moniliform 178
Nakh 317 Oak 80 Palaquium 375, 29
Monimiaceae 101
Name Proposal 87 Oats 520 Palea 519
Monkey flower 407
Names of Cultivated Plants 87 Obcordate 183 Palea 184
Monkey fruit 455
Names of Different Taxa 88 Obdiplostemonous 169, 201 Paliurus 286
Monochlamydae 72
Napiform 179, 243 Obdiplostemony 250 Palmaceae 75
Monochlamydeae 25
Naravelia 221 Obtuse 183 Palmae 26, 98, 500, 504, 509, 546
Monocots 40
Narcissus 106, 477, 478 Ochlandra 523 Palmae1 469
Monocotyledoneae) 13
Narcissus) 8 Ochroma 269, 420 Palmales 31
Monocotyledones 20, 455, 456, 469
Nardus 520, 524 Ocimum 419, 213, 551 Palmarosa oil 523
Monocotyledons 72, 545
Narenga 523 Odontoglossum 470, 474 Palmately Compound Leaf 189
Monocotyledons) 12
Nargis 479 Oenothera 119 Palwal 331
Monodora 218
Nariyal 502 Offset 181 Palynology 106
Monograph 148
Nasturtium 97 Oil palm 500 Pampas Grass 523
Monomials 88
Natural Classifications 20 Okra 260 Panama rubber 455
Monopetalae 23
Natural classification systems 22 Olacales 25, 231 Panax 346, 306, 550
Monophyletic 140
Natural system 23 Olacineae 25 Pancratium 477, 479
Monophyletic origin of angiosperms
Natural System Approach 10 Oldenlandia 349 Pandanaceae 29, 470, 504
140
Natural system of classification 11 Olea 378 Pandanales 28
Monosulcate pollen grains 107
Natural systems 19 Oleaceae 25, 543 Pandanus 143
Monsonia 271
Nauclea 350 Omphalodes 394 Pandorea 409
Monstera 507
Navia 480, 482 Omphalogramma 371 Panic grass 518
Montia 254
Neem tree 282 Onagraceae 102 Panicle 194
Mooli 244
Nelumbaceae 229 Oncidium 470, 472, 474 Panicum 518, 522, 523
Moongphali 309
Nelumbo 227, 292, 547 Onion 493 Pansies 247, 251
Moraceae 29, 545
Nelumbonaceae 109, 299, 429 Onosis 119 Papaver 22, 27, 528, 548
Moraceae1 429
Nemacladus 365 Onosma 394 Papaveraceae 24, 213, 300, 528, 540
Moraceae, Urticaceae 34
Nemesia 407 Operational Taxonomic Units 116 Papaveraceae, Fumariaceae 34
Morinda 350
Neo-Adansonian principles 115 Ophioglossum 104 Papaverales 233
Moringaceae 102
Neo-Darwinism 50 Opium 231 Paphiopedalum 474
Morning glory 396
Neottia 470, 471 Opium poppy 231 Papilionaceae 74, 32, 542
Morphotaxa 85
Neotype 86 Opium poppy 233 Papilionaceous 307
Morus 454, 551
Nepenthaceae 25, 399, 400 Opposite 186 Papilionoideae 306, 76
Mosses 22
Nepeta 419 Opposite decussate 186 Papilionoideae 299, 531
Moth 309
Index 561
Pappus 196, 357 Periwrinkle 387 Piperaceae1 429 Polyscias 348

Papyrus plant 517 Pernettya 367 Piperales 28 Polysepalous 196
Parallel 191 Peronema 423 Pistachio-nuts 293, 296 Pomaderis 285
Para-rubber 446 Personales 25 Pistacia 293, 324 Pome 209
Parasite 177 Petals 169 Pistia 507, 508 Pomegranate 329
Paratype 87 Petha-kaddoo 331 Pistil 65 Poncirus 280
Parietal 205 Petrea 425 Pistillode 203 Pongamia 309
Parietales 24, 337, 124, 99 Petrocoptis 251 Pisum 306 Pontederiaceae 469
Parietaria 451, 329 Petroselinum 343 Pitcairnia 480, 482 Poplar 463
Paripinnate 190 Petunia 399, 363 Pithecellobium 302 Populus 109
Paris 490, 29, 492, 493 Peucedanum 342 Pit-papra 238 Porana 395
Paris Code 81 Pflanzenfamilien 12 Piyaz 493 Portable Freezer Unit 58
Parishia 296 Phaeophyceae 28 Place of the Origin of Angiosperms Portable refrigerator 58
Parkia 299, 29 Phalaenopsis 474 144 Portulaca 253, 548
Parkinsonia 303 Phalaris 522, 523 Plagiobothrys 392 Portulacaceae 25, 253, 444
Parnassia 111, 104 Phalsa 268 Plagiopteron 266 Potamogetonaceae 85
Parnassiaceae 111 Phanerogamae 27 Plantae 47 Potato 399
Paronychia 250, 108 Phanerogams 8 Plantaginaceae 30, 544 Potenti11a 317
Parseley 343 Pharus 524 Plantaginales 30 Potentilla 312, 549
Parsnip 343 Phaseolus 9, 136 Plantago 426 Poterium 312, 527
Parsonsia 385 Phellodendron 280 Plantain 426 Pothos 507, 508
Parthenium 99 Phenetic taxonomy 11 Plantanaceae 429 Pouzolzia 451
Parthenocissus 287, 110 Phenons 118 Plant Chemotaxonomy 121 Precipitin Reaction 135
Parts used as Epithets 90 Philadelphus 318 Plant collection 55 Precipitins 135
Paspalum 518, 523 Phillyrea 380 Plant Identification 69 Prefixes of numbers 89
Passiflora 336, 406, 407 Philodendron 507 Plant Nomenclature 78 Premna 425
Passifloraceae 102, 371, 542 Philyderaceae 469 Platycodon 363 Prestonia 385
Passiflorales 25 Phlomis 100 Platystemon 233 Prickly Poppy 233
Passion fruit 338 Phlox 401 Pleiochasium 194 Primitive Angiosperms 143
Pastinaca 343 Phoenix 500, 501, 502, 503 Pleurothallis 472 Primrose 371
Paullinia 290, 406 Phool Gobhi 244 Plum 312 Primula 21
Paulownia 406, 285 Phool Matar 310 Plumbaginaceae 29, 438, 543 Primulaceae 25, 543
Pavonia 257 Phoont 331 Plumbaginales 29 Primulales 25
Payena 377 Phoradendron 443 Plumbago 372 Principes 28
Pea 309 Phormium 493 Plumeria 384 Principle of Priority 86
Peach 312 Phylica 285 Pneumatophorous 179 Principles of Phylocode 92
Peanut 309 Phyllanthus 323 Poa 103, 518, 523 Prionium 497
Peanut butter 308 Phyllaries 356 Poaceae 52, 518, 524, 539, 546 Priority 81
Pear 317 Phylloclade 181 Podina 421 Pritchardia 502
Pearl 522 Phyllode 181 Podophyllaceae 109 Probable Ancestors of Angiosperms
Pedaliaceae 25, 544 Phyllostachys 523 Podophyllum 109 141
Pedalium 412 Phyllyrea 101 Podostemaceae 25, 327 Problems of Hierarchy 47
Pedicel 195 Phylocode 91 Podostemonaceae 29 Prodromus 23
Pedicularis 405, 382 Phylogenetic Classifications 20, 39 Podostemonales 29 Pronuba 492
Peeli Kaner 387 Phylogenetic system 11, 30 Pogostemon 421 Prophyll 184
Peepal 455 Phylogenetic tree 128 Poikilospermum 452 Prop or Stilt 179
Peganum 111, 123, 549 Phylogeny 138 Poinsettia 446 Prosopis 299
Pelargonium 271 Phylogeny of angiosperms 138 Poinsettia 449 Proteaceae 26
Pelletiera 369 Physalis 399, 443 Poison hemlock 121, 464 Proteales 29
Pellionia 453 Physaria 106 Polanisia 86, 463 Proteins 126
Penaeaceae 429, 416 Physocarpus 315 Polemoniaceae 103, 544 Protista 47
Pennisetum 518, 522, 524 Physopsis 423 Polemoniales 25 Prunus 10
Penstemon 405 Physostigma 310 Polemonium 401 Pseudotsuga 124
Pentace 266 Phytelephas 500, 502, 503 Polianthes 479 Psidium 323
Pentapetes 263 Phyteuma 362, 280 Pollen morphology 108 Psilotum 122
Pentaphragma 111 Phytolacaceae 428 Pollinia 472 Psittacanthus 443
Pentaphragmataceae 111 Phytolacca 109 Pollinium 381 Psophocarpus 309
Pentas 350 Phytolaccaceae 109, 249, 314 Polyadephous 201 Psoralea 306, 549
Pentoxylon Theory 143 Piazi 494 Polyalthia 218 Psychotria 349
Peperomia 441 Picrorhiza 407 Polycarpaea 252 Ptelea 277
Peplis 328 Pieris 367 Polyclave 71 Pteridophyta 27
Pepo 209 Pigeon pea 309 Polyclave identification 71 Pteridosperms 12, 39
Perennial 177 Pilea 451 Polyclave-type devices 69 Pteridosperm Theory 142
Pereskia 339 Pilkhan 455 Polygaleae 24 Pterisanthes 287
Perfectae 20 Pilocarpus 280, 549 Polygalineae 24 Pterocarpus 307
Pergularia 384 Pimenta 324, 426 Polygonaceae 25, 435, 536, 544 Pteropetalum 240
Perianth 64 Pimpinella 342 Polygonaceae1 428 Pterospermum 262
Perianth 195 Pina-cloth 480 Polygonales 29 Pubescent 181
Perigynium 516 Pinax Theatri Botanici 10, 79 Polygonum 435, 537, 551 Pulmonaria 394
Perigynous 22 Pind Khajoor 503 Polymorphosporophyta 108 Pulsatilla 223
Perigynous 195, 213 Pineapple 480 Polypetalae 23 Pulses 308, 309
Perilla 421 Pinetum 162 Polypetalous 197 Punched cards keys 70
Period of Linnaeus 10 Pinnately Compound Leaf 190 Polyphyletic 140 Punica 327
Periploca 381, 402 Pinus 100, 299, 300 Polyphyletic Origin 140 Punicaceae 85
Peristrophe 414 Piper 441, 503 Polyploid 104 Pupalia 432
Peristrophe 415 Piperaceae 25, 428, 544 Polyploidy 493 Puya 480
562 Index
Pycreus 515 Rheum 435, 551 Sacred lotus 229 Scilla 493
Pygeum 312, 532 Rhipidium 399 Sadabahar 387 Scindapsus 509
Pyrethrum 359 Rhipsalis 339, 277 Safed sarson 245 Scirpus 21, 513, 514, 517, 518
Pyrolaceae 365 Rhizobium 307 Safflower 359 Scitamineae 28, 482, 485, 490
Pyrus 80 Rhizome 181 Saffron 475 Scitamineae1 469
Rhizophoraceae 323 Sage 419 Scleria 514, 518
Quadrinomials 89
Rhodea 492 Sageretia 286 Scolyopus 491
Quamoclit 395
Rhododendron 100, 447, 448, 449 Sagina 250 Scoparia 406, 283
Quercus 80
Rhodophyceae 28 Sagittaria 510, 512, 513 Scrophhulariales 32
Quillaja 317
Rhoeadales 29 Sagittate 187 Scrophularia 97, 267
Quince 317
Rhoeo 494, 495 Sago 503 Scrophulariaceae 25, 535, 544
Quincuncial 197
Rhoicissus 288 Sago palm 500, 503 Scrophulariales 32
Quinine 350
Rhomboid 187 Salicaceae 12, 111, 355, 545 Scutellaria 419
Quisqualis 321
Rhubarb 435 Salicaceae1 429 Scytopetalaceae 257
Raceme 194 Rhus 293 Salicales 28 Seb 316
Racemose 192 Rhynchodia 387 Salicornia 429, 354 Secale 518, 521
Rachilla 184, 519 Rhynchospora 514, 517, 518 Salix 109, 400, 551 Sechium 331
Rachis 185 Ribbon grass 523 Salmalia 269 Secondary key characters 70
Radermachera 411 Ribes 318 Salomus 369 Sedge 514
Radical 185 Rice 518 Salpiglosis 399 Seed 67, 176
Radical or basal 186 Rice 521 Salpiglossidaceae 401 Seed ferns 12, 413
Radish 244 Rice-paper plant 348 Salpiglossis 136, 30 Selaginaceae 417
Ragi 522 Ricinocarpos 446 Salsola 429, 290 Selago 405
Rai 245 Ricinus 446, 551 Salvadoraceae 34, 47 Sem 309
Rajmah 309 Rinorea 247, 421 Salvia 419, 214 Semantides 122
Ramie 453 Ritha 292 Samanea 302 Semecarpus 293
Ramphal 219 Rivinia 109 Samara 210 Semibegoniella 334
Ranales 24, 29, 40 Rizka 309 Sambucaceae 355 Semi-inferior 204
Ranalian concept of evolution 31 Robinia 307 Sambucus 353, 29, 297 Senebiera 244
Randia 350, 30 Rochester Code 81 Sandoricum 282 Senecio 356
Rank Alteration 87 Romulea 475 Sanguinaria 232, 319, 548 Sengri 245
Ranks and Endings of Taxa 86 Rondeletia 349 Sanguinarine 233 Senna 303
Ranks of Plant Classification 19, 244, Root 62, 176 Sani 310 Sensitive plant 299, 264
145 Rorippa 243 Sanicula 342 Sepals 169
Ranks of Taxa 5 Rosa 22, 532, 549 Sansevieria 490, 493 Series of ranks and endings 5
Ranunculaceae 24, 221, 513, 526, 527, Rosaceae 25, 407, 527, 531, 532, 542 Santalaceae 26, 348 Serjania 290
540 Rosaceae 312 Santanales 29 Serological Reactions 135
Ranunculidae 37 Rosales 25 Sapindaceae 25, 292, 531, 541 Serology 134
Ranunculus 22, 400, 547 Roscoea 488, 490 Sapindales 25 Serotaxonomy 134, 135
Rapatiaceae 469 Rose 312 Sapindus 289 Sesame 412
Raphanus 243, 385 Rosidae 38 Sapium 446 Sesame oil 412
Raphia 500, 503 Rosmarinus 419, 435 Sapodilla 375 Sesamum 412
Raspberry 163, 312, 317 Rostellum 472 Sapodilla plum 376 Sesbania 307
Rattan 503 Rostkovia 497 Saponaria 250, 548 Sessile 185
Ratti 309 Rotala 328 Sapotaceae 25, 543 Setaria 518, 522
Rattlebox 306 Rotund 187 Saraca 303, 341 Sexuality in plants. 9
Rauvolfia 384, 550 Round gourd 331 Sarcandra 101 Sgsv 59
Ravenala 485, 486, 487 Roxburghiaceae 469 Sarchochlamys 451 Shahtoot 455
Ray florets 356 Royal Botanical Garden 160 Sarcobatus 430 Shakarkandi 395
Rebutia 340 Royal palm 500, 502 Sarcostemma 384 Shaljam 244
Red water-lily 228 Royal water-lily 228 Sarpgandha 387 Shantra 279
Reevesia 265 Roydsia 240 Sarraceniaceae 231 Shape of Leaf 187
Reference 149 Roystonea 500, 502 Sarraceniales 29 Sharifa 218
Reference Book 149 Rubber 449, 450 Sarson 245 Shepherd’s purse 245
Reference Reaction 135 Rubber plant 454 Satice 374 Shiajeera 343
Reflexa 397 Rubia 349 Saunf 343 Shikakai 302
Regma 210 Rubiaceae 25, 349, 533, 534 Sauromatum 509 Shisham 306
RegNum 93 Rubiales 25 Saxifraga 318 Shrub 177
Rejection of Names 88 Rubus 312 Saxifragaceae 106, 172, 532, 542 Shrubs 20
Relationship 374 Rudraksh 265 Scanning electron microscopy 113 Sibling species 50
Relationship tree 128 Ruellia 414 Scape 182 Sicydium 330
Remusatia 509 Rules and Recommendations of ICBN Schefflera 346 Sida 257
Reniform 187 82 Scheuchzeriaceae 103, 251 Sideroxylon 377
Replacement names 84 Rumex 21, 551 Schinopsis 296 Silene 250
Replum 203 Runner 182 Schinus 293 Silica bodies 98
Resedaceae 231 Ruscus 490, 491, 492, 493 Schizandra 137 Silicula 210
Respiratory 179 Russelia 406 Schizandraceae 137, 405 Siliqua 210
Restionaceae 470, 498, 524 Ruta 276 Schizanthus 399, 245, 260 Silybium 358
Resupination 472 Rutaceae 25, 276, 530, 541 Schizocarp 343 Simaroubaceae 280
Retention of Names 88 Rye 521 Schizopepon 330, 532 Simarubaceae 284
Reticulate 191 Schizophyta 28 Simple Leaf 191
Sabal 500, 502, 503
Retinaculae 381 Schleichera 289 Siphocampylus 362
Sabiaceae 25, 289
Rhamnaceae 25, 285, 309, 541 Schlumbergera 340 Siris 302
Sabudana 503
Rhamnales 29 Schoenus 514 Sisymbrella 97
Saccharum 79, 518, 522, 523, 524
Rhamnus 285, 549 Schrankia 299 Sisymbrium 243
Sacred lotus 227
Rhaphidophora 507 Scientific names 79 Sisyrinchium 475
Index 563
Sitaphal 331 Stipa 518, 523, 524 Tapioca 449 Tradescantia 494, 495
Sitopsis 106 St. Louis Code 82 Tap Root 177 Tragia 446
Skimmia 277 Stolon 182 Taramira 245 Tragopogon 123
Smilacaceae 494 Stomata 98 Taraxacum 359 Transitional phylogenetic system 27
Smilax 490, 491, 492, 493, 494, 551 Stramonium 400 Tarbooj 331 Translators 381
Snake gourd 331 Strawberry 317 Tautonym 87 Transmission electron microscopy 113
Snapdragon 405 Streblus 456 Taxaceae 111 Trapa 102, 450
Soapnut tree 292 Strelitzia 485, 486, 487 Taximetrics 115 Traveller’s joy 223
Solanaceae 25, 310, 534, 543 Strelitziaceae 486 Taxon 78, 310 Traveller’s Tree 486
Solanaceae 397 Streptolirion 494, 495 Taxonomic characters 46 Tree 177
Solanaceae, Convolvulaceae 34 Strobilanthes 414 Taxonomic works of India 14 TreeBASE 93
Solanum 101 Strophanthus 385 Taxonomist 3 Tree of life 91
Soldanella 369 Struthanthus 443 Taxonomy 1, 411 Trees 20
Soleirolia 453 Strychnos 124, 550 Teak 425 Trewia 450
Sonchus 355 Stylidaceae 355 Techniques used in molecular taxonomy Tribes 52
Sonneratia 328 Stylidiaceae 98 129 Trichodesma 392
Sonneratiaceae 327 Stylopodium 342 Tecoma 409 Trichomes 99
Sophora 307 Styraceae 375 Tecomaria 409 Trichopodiaceae 102
Sorbaria 315 Suaeda 429 Tectona 423 Trichopus 102
Sorbus 312 Subforms 48 Tejpat 279 Trichosanthes 330
Sorghum 518, 522 Subspecies 47 Tendril 182, 330 Tricolpate pollen grains 107
Sorosis 210, 455 Subspecies 48 Tepals 64 Trientalis 369
Sorrel 437 Subterranean 182 Tephrosia 310 Trifolia 22
Sour cherry 317 Subulate 187 Teramnus 310, 549 Trifolium 97
Sowa 343 Subvarieties 48 Terminalia 321, 260, 549 Trigonella 307
Soya 309 Sucker 182 Terpenoids 123 Trimorphosporophyta 108
Soybean 308, 309 Sugarbeet 429 Tessarandra 378 Trinomials 89
Soymida 282, 387 Sugarcane 518, 522 Tetracentraceae 101 Triosteum 353
Spadix 194 Sugar Maple 290 Tetracentron 101 Triphala 323
Sparganiaceae 99 Sukhdarsan 478 Tetractomia 277 Triphasia 277
Sparmannia 265, 430 Sumac 293 Tetradynamous 202, 243 Triplaris 435, 536
Spathicarpa 508 Sunflower 359 Tetrameranthus 218 Tristania 324
Spathiflorae 28 Sunnhemp 310 Tetrapanax 348 Triticum 105, 518, 520, 521, 522, 523,
Spathiphyllum 508, 509 Supari 503 Tetrapoma 244 524
Spathodea 411 Superior 204 Tetrastigma 287 Tritonia 475
Spatholobus 309 Surajmukhi 359 Teucrium 419 Triumfetta 265, 548
Spathulate 187 Sushrut Samhita 13 Thalamiflorae 24 Triuridaceae 28, 470
Speciation 130 Sutera 405 Thalictrum 105, 385, 387 Triuridales 28
Species 19 Sweet basil 421 Thallophyta 27 Trochodendraceae 101
Species Plantarum 10, 21 Sweet Basil 422 Theaceae 100 Trochodendron 101
Specific epithet 47, 80 Sweet cherry 317 The Linnaean Society of London, 10 Trollius 221
Specularia 363 Sweet pea 309, 306 Theobroma 80 Tuber 182
Spergula 250, 548 Sweet potato 395 Theophrasta 80 Tuberous 179
Spergularia 250 Swietenia 281 Theophrastaceae 369 Tubiflorae 30
Sphaeranthus 359, 550 Syconus 210 Theory of polyphylesis 140 Tulipa 490, 491, 492, 493
Spigelia 389 Symbegonia 334 Thespesia 258, 415 Tulsi 421
Spike 194 Symphoricarpos 353 Thevetia 384, 345, 550 Tunt 455
Spikelet 194, 519 Symphytum 394 Thladiantha 330 Turmeric 489
Spilanthes 359 Symplocaceae 375 Thorn 182 Turnip 243
Spinach 429 Sympyandra 363 Thrinax 500 Turpinia 290
Spinach 430 Synadenium 450, 551 Thunbergia 414, 421 Tussilago 359
Spinacia 429 Synandrous 201 Thymeleaceae 429 Twenty four principles 33
Spine 182 Synanthae 28 Thymus 419, 208, 209, 551 Twinner 182
Spiraea 312 Synaptantha 350 Thyrse 194 Twisted 197
Spirodella 123 Syncarpous 204 Thysanolaena 523 Tylophora 380, 550
Spondias 293 Syngenesious 202 Til 412 Type 81
Sporobolus 520, 523 Synonym 87 Tilia 265, 266 Type Method 86
Squamellae 385 Syntype 87 Tiliaceae 25, 306, 529, 541 Types of aestivation 197
Stachys 419 Syringa 378 Tiliaceae, Sterculiaceae, Bombacaceae Types of androecium 200
Stachytarpheta 423 Systema Naturae 21 34 Types of corolla 198
Stamen 8, 21 Systematics 1, 2, 3 Tillandsia 479, 480, 482 Types of fruit 207
Staminode 169, 201 Systems of Classification 20 Time of Angiosperms Origin 141 Types of inflorescence 193
Standard 307 Syzygium 323, 549 Time of great herbalists 9 Types of leaf 190
Stapelia 381 Tinda 331 Types of stipule 189
Tabebuia 409, 411
Staurogyne 414 Tobacco 399 Types of Stipule 187
Tabernaemontana 384, 385
Stelechocarpus 218 Toddalia 277, 549 Type specimen 86
Taccaceae 469, 482
Stellaria 250, 548 Toddy 503 Typha 22, 504, 505, 506
Taeniantherum 126
Stem 63 Tomato 400 Typhaceae 26, 504, 506, 546
Tagetus 359, 360
Stenocereus 340 Toona 281 Typhaceae1 470
Talauma 214, 255
Stenopalynous 107 Topotype 87 Typhonium 507, 508
Talinum 253, 2
Stephanotis 382 Torenia 407 Typhostemma 336
Tamarind 304
Sterculia 262 Toria 245 Typological Concept of Species 49
Tamarindus 303, 118, 73
Sterculiaceae 25, 541, 262 Tournefortia 392
Tamariscineae 249 Ulmaceae 29
Stereospermum 411 Toxicodendron 297
Tamatar 400 Ulmus 21, 97
Steroids 124 Trachymene 342
Tanacetum 359, 550 Umbel 194, 342
Stigma 204 Trachyspermum 343
564 Index
Umbellales 25, 34, 341 Vasculum 56 Viscaceae 105 Xanthorrhoea 491, 493
Umbelliferae 25, 29, 341, 342, 345, 348, Vegetables 309 Viscoideae 110 Xanthorrhoeaceae 100
533, 542 Vellozia 100 Viscum 443 Xanthosoma 509
Uncaria 349 Velloziaceae 100 Vitaceae 29, 286, 541 Xanthoxylum 80
Uncinia 514, 518 Venation 191 Vitex 423 Xantolis 377
Unicostate 191 Ventilago 285, 424 Vitis 286 Xerophyte 177
Unifoliate 189 Veratrum 493, 551 Vivania 273 Xylia 299
Uniola 519, 524 Verbascum 405, 227, 228 Voucher 149 Xylocarpus 281
Unipalynous 107 Verbena 423, 405, 406 Vriesea 480, 482 Xylopia 218
Unipinnate 190 Verbenaceae 25, 72, 306, 307, 309, 423, Vrikshayurveda 9, 13 Xyridaceae 101, 469
Unisexuales 26, 260 535, 544
Wahlenbergia 362 Ylang-ylang 218
Unona 219 Vernacular 79
Walsura 281, 101, 549 Yucca 105, 490, 491, 492, 493
Upas tree 456 Vernonia 99
Waltheria 263
Uraria 219, 547 Veronica 21, 248, 249 Zanonia 331
Water grass 504
Urd 309 Versatile 202 Zanthorhiza 105
Water hemlock 343
Urena 260, 451, 548 Verticillaster 28, 192, 194 Zanthoxylum 276, 97, 100, 549
Watermelon 331
Urera 451 Vervain 425 Zardalu 316
Water plaintain 510
Urginea 493 Vetiveria 523 Zea 518, 519, 521, 522, 523
Waxes 125
Urtica 22 Vexillary 197 Zebrina 494, 495
Wax palm 500
Urticaceae 26, 451, 545 Vexillum 307 Zedoary 490
Wheat 518
Urticaceae1 429 Viburnum 352 Zeera 343
Wheat 521
Urticales 29 Vicia 126, 385, 387, 284 Zephyranthes 477, 479
Wheat grass 518
Used 97 Victoria 162 Zeuxine 470, 472, 473
Whorled 186
Usteria 389 Viena Code 84 Zimikand 509
Wiegela 352
Utricle 210 Vienna Code 81 Zingiber 488, 489, 551
Wielandia 447
Vienna Rules 84 Zingiberaceae 76, 483, 488, 490, 537,
Vacciniaceae 368 Wild date palm 503
Vigna 136 546
Vaccinium 366, 368 Willow 463
Vilayati kikar 302, 304 Zingiberidae 39
Vahila 247 Wine palm 502
Vilayati Sem 309 Zinnia 119
Valerianaceae 349, 30, 360 Winteraceae 98
Vinca 384 Zizyphus 285
Validly published 88 Wisneria 510
Vine 177 Zornia 307
Vallaris 387 Wisteria 309
Viola 247, 548 Zoysia 524
Valvate 197 Withania 399, 550
Violaceae 24, 540 Zygogynum 215
Vanda 470, 471, 474 Woodfordia 327
VIOLACEAE 246 zygomorphic 169
Vanilla 470, 473, 474 World’s largest herbarium 152
Violales 249 Zygomorphic 196
Varieties 47 Wrightia 387
Violas 247 Zygophyllaceae 111
Variety 48
Violets 247 Xanthium 360
Vasaka 415
Virgins-bower 223 Xanthorhiza 221
Vasculares 23

Plant Taxonomy

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Plant Taxonomy

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PLANT TAXONOMY

Tata McGraw-Hill Education Private Limited

Copyright © 2009 by Tata McGraw-Hill Publishing Company Limited

This edition can be exported from India only by the publishers,

ISBN (13): 978-0-07-014159-9

Managing Director: Ajay Shukla

Cover Printer: SDR Printers

The McGraw-Hill Companies

3.3 Types of Systems of Classification 19

7.3 Identification with Keys 70

9.5 Palynology in Relation to Taxonomy 106

Test Your Understanding 133

16. Herbarium 152

20.3 Root 177

21.21 Bombacaceae (Bombax Family) 268

21.57 Asterales 355

21.93 Loranthaceae (Mistletoe Family) 443

Table A.6 Papaveraceae and Cruciferae (Brassicaceae) 528

Jun 2009 O P Sharma

May, 1993 O P Sharma

1.2 SYSTEMATICS AND THE SYSTEMATIST

6. As an observer, a taxonomist observes the characteristics of organisms, population and taxa

1.2.3 Every Human Being is a Taxonomist

1.3 OBJECTIVES, GOALS AND AIMS OF PLANT SYSTEMATICS

3. To be aware of the importance of taxonomic relationships in plant systematics.

1.3.3 Aims for the Study of Regional Flora

1.4 HIERARCHICAL STAGES OF A SYSTEMATIST

1.5 HIERARCHICAL CATEGORIES OF TAXONOMY

1.6 BASIC COMPONENTS OF TAXONOMY

Ranks of Taxa Endings of ranks above genus Examples

Division phyta Pterophyta, Magnoliophyta

1.7 FUTURE OF PLANT TAXONOMY

Test Your Understanding

2.1.2 Age of Theophrastus, Secundus, Dioscorides and Parasara

2.1.3 Taxonomy in Middle Ages

2.1.5 Taxonomy during Seventeenth Century

2.1.6 Period of Linnaeus

2.1.7 Natural System Approach

2.1.8 The Phylogenetic Approach

2.1.9 Some Current Contributions

2.2 TAXONOMY OUR CONTEMPORARY

2.3 CHRONOLOGICAL DEVELOPMENT OF TAXONOMY IN INDIA

2.4 SOME INDIAN JOURNALS RELATED TO TAXONOMY

3. Bulletin of the Botanical Survey of India

Table 2.1 Chronology of the major published taxonomic works of India

Year A.D. Researcher/Explorer Major published/botanical work

1565 Garcia d’Orta Published Os Colquios in Goa; described common

1903 D. Prain Bengal Plants

1977 M. Oomachan The Flora of Bhopal

2.5 A NOTE ON BOTANICAL SURVEY OF INDIA

Test Your Understanding

3.2 RANKS OF PLANT CLASSIFICATION

3.3 TYPES OF SYSTEMS OF CLASSIFICATION

3.4 SOME IMPORTANT SYSTEMS OF CLASSIFICATION

Imperfectae (Flowerless plants)

I. Herbae (herbs) Dicotyledones

Perfectae (Flowering plants)

II. Arborae (shrubs and trees)

3.4.2 Carl Linnaeus1 (1707–1778)

8. Octandria (with 8 stamens), e.g. Fagopyrum.

3.4.3 Antoine-Laurent de Jussieu (1748–1836)1

Stamina perigyna (stamens perigynous)..............................................................................VI

3.4.4 Augustin Pyramus de Candolle (1778–1841)1